The gobiid fishes of temperate Macaronesia (eastern Atlantic)

J. Zool., Lond ( 1 984) 204, 363-4 12

The gobiid fishes of temperate Macaronesia (eastern Atlantic)

(With 3 plates and 17 figures in the text)

Eight species of gobiid tishes have been recorded from the islands comprising temperate Macaronesia (Azores. Canaries. Madeira and Salvages). These are (;ohiir.\ n/,ycr L.. G ui(ru/io Risso, G. ,yrr s r c ~ w ~ i i Mi I ler. G /iuyuiie,//ir 5 L.. L~,\irc,ir,r,y/:o/iiirs / w / w ) / i \cru/i/.\ Maul. .MuirligohIir.\ r ~ ~ u d c ~ r c ~ m i . ~ (Valenciennes) and 7'/ioro,yohirr\ c~p/iippirrrir\ (Lowe). with L. . \ i w i i

(Risso) remaining unconfirmed. The identification, records, characteristics. and habitat of these species in the Macaronesian islands are considered. T h e genus . l / ~ / d i , y d i i i i . ~ Miller is fully defined and the type-species. Gohiir, rnrcikc~rcw\i.\ Valenciennes, endemic to Madeira and the Canaries. described in detail. The transfer of the little-known West African Gohiir, nigri Giinther to .\/uir/igohirt.s is suggested. Phylogenetic discussion of.llair/i,y.ohiir\. in relation to Atlantic-Mediterranean and Ponto-Caspian forms. indicates its position as the sister group to Gohim. The modern Macaronesian gobiid Fauna seems to be derived from the warm-temperate eastern Atlantic-Mediterranean marine province. The presumed absence of .If. /iiurkw/i.ri.\ from the Azores. and the pattern of meristic variation i n Gohiir\ pu~yumd/u.\, support the view that the continental rise islands d o not have a direct Faunal interrelationship with the Azores. and that common possession of Atlantic-Mediterranean species reflects supply from the same outside source rather than by interchange. The likelihood of finding more gobiid species in Macaronesia is noted. A model of speciation in Muii/igo/>rir.\, in relation to hydrographic events. is suggested.

Appendix I summarizes features of marking and head lateral-line system useful for distinguishing juvenile G. pugurw//ir.\ and .\I. r i inc lrw/ i . \ i v . Appendix I I provides a redescription of the holotypc of Gohiiis ~ i g r i , and a discussion of geographical occurrence. habitat. and affinities for this species.

Contents

Introduction . . . . . . . . . . . . . . . . . . . . Abbreviations . . . . . . . . . . . . . . . . . . . . Systematic list . . . . . . . . . . . . . . . . . . . . The genus .Iluii/igohrir.\ Miller 198 I . . . . . . . . . . . . . . Origin and relationships of the Macaronesian gobiid fauna . . . . . . References . . . . . . . . . . . . . . . . . . . . Appendices . . . . . . . . . . . . . . . . . . . .

Page 363 365 365 384 397 402 406

Introduction

Macaronesia is a term used in phytogeography (Good, 1953) for a floristic region situated not, as might be supposed, in the South Pacific, but embracing the Atlantic islands of Madeira, the Canaries, and the Cape Verdes, off the coast of north-west Africa, as well as

363 0022-5460/84/110363+50503.00/0 0 1984 T h e Zoological Society of London

364 P. J . MILLER

the mid-oceanic Azores (Fig. l(a)). The more northern groups above the Tropic of Cancer, comprising Madeira, the Canaries, and the Azores (geologically younger than Cape Verdes), have a coastal marine fauna in which Lusitanian-Mediterranean species predominate. These islands are thus recognized as outlying provinces ofthe continental Lusitanian region (Briggs, 1974) and, for them, "temperate Macaronesia" may be adopted in zoogeography as a useful collective name (Fig. l(b)). Although of similar, if much earlier, geological formation, the Cape Verde islands, far to the south, have an essentially tropical West African marine fauna (Briggs, 1974) and are excluded from the present study.

40 - . ...yes

Madeira

Islands :"' *

20 O N .

... Cape Verde Islands

Madeira a b

** D 30 O N

Canary 0 od 0 Islands G

H 1 I\ I

1 5 O W 20 l O 0 W

FIG. I . (a) Islands of Macaronesia in relation to Europe and Africa. (b) Temperate Macaronesia: islands mentioned in text and Table VI. Canary Islands: C , Gran Canaria; F, Fuerteventura; G, Gomera: H, Hierro; L, Lanzarote: LP, La Palma; T, Tenerife. Madeira: D, Desertas; Azores; Co, Corvo: Fa, Fayal; FI, Flores; Gr, Craciosa, SJ, SBo Jorge; SM, SBo Miguel; SMa, Santa Maria; P, Pico; Te, Terceira, SG, Straits ofGibraltar.

The northernmost Macaronesian islands on the African continental rise are Madeira and neighbouring islets (Funchal: 32'38 'N, 16'54 'W), about 400 miles west of Morocco, on the south end of the Madeira-Tore Rise. The Canaries include seven major islands along the broad Canary plateau, extending over a sea area of about 300 miles E-W by 120 miles N-S (27'37 'N, 17'57 'W to 29'25 'N, 13'29 'W), with Fuerteventura only 65 miles from Cape Juby, Morocco. Two clusters of small islands and reefs, the Salvage Islands (Selvagems) (30'2'N, 16'2'W), lie between Madeira and the Canaries. Truly oceanic in position, about 900-1200 miles west of Lisbon, the Azores archipelago, of nine major islands, stretches for about 300 miles E-W, by 130 miles N-S, across the Mid-Atlantic ridge, from 36'55 ' N and 25'1 'W to 39'43 ' N and 3 1'7 'W, on the western end ofthe East Azores fracture zone. Along the line of the Azores and that of the continental rise islands, scattered seamounts, some formerly exposed, converge towards the Cadiz Sea off southern Iberia (Uchupi et al., 1976).

The present paper is based on various collections of gobioid fishes from Madeira, the Canaries, and the Azores. A systematic list reviews identification of species in the light of modern work on Atlantic-Mediterranean gobiid systematics and literature relating to

GOBllD FISHES O F TEMPERATE MACARONESIA 365

temperate Macaronesian fishes. In particular, special attention is paid to an endemic goby, originally described from Madeira as Gohius maderensis by Valenciennes ( 1 837). Examin- ation of the types, and much recent material, has now permitted a reappraisal of the species, with definition of a new genus (Mauligohius Miller, I98 1 ) and consideration of its systematic affinities. The origin and relationships of the temperate Macaronesian gobies are discussed with reference to hydrography and climatic changes.

Abbreviations

Collections. BMNH: British Museum (Natural History); CANCAP-RMNH: CANCAP- project, Rijksmuseum van Natuurlijke Historie, Leiden (den Hartog & Lavaleye, 198 I ) ; HUJ: Hebrew University, Jerusalem; MHNG: Museum d’Histoire Naturelle, Geneva; MMF: Museu Municipal do Funchal, Madeira; MNHN: Museum National d’Histoire Naturelle, Paris; NHMW: Naturhistorisches Museum, Vienna: ZMUC: Universitetets Zoologiske Museum Kobenhavn.

Meristics. A: anal fin; C: caudal fin; D1, D2: first and second dorsal fins; LL: scales in lateral series; P: pectoral fin: PD: predorsal scales: TR: scales in transverse series: V: pelvic disc; VC: vertebrae.

Morphometrics (Tables I , IV). Ab: anal fin base; Ad and Aw: body depth and width at anal fin origin; C1: caudal fin length; CHd: cheek depth: CP and CPd: caudal peduncle length and depth; D1 b and D2b: first and second dorsal fin bases: E: eye diameter; H and Hw: head length and width; I: interorbital width: PI: pectoral fin length; PO: postorbital length; SL: standard length; SN: snout length: SN/A and SN/AN: distance from snout to vertical of anal fin origin and anus; SN/DI and SN/D2: distance from snout to origin of first and second dorsal fins: SN/V: distance from snout to vertical of pelvic disc origin; V/AN: distance from pelvic disc origin to anus; Vd: body depth at pelvic disc origin: V1: pelvic disc length.

Lateral-line system (Tables 11, V). Series: AD, anterior dorsal; OP, opercular: OS, oculoscapular; PM, preopercular-mandibular; PO, postorbital; SO, suborbital; terminology of rows and canal-pores as Miller (In press), based on Sanzo ( 19 1 I ) .

Systematic list

The order of species known from Macaronesia follows the essentially alphabetical sequence of Miller ( 1 9 7 3 ~ ) . Synonymy is restricted to citation of works referring to Macaronesian or adjacent areas; full synonymies are given by Miller ( 1 9 7 3 ~ ) . Description is limited to a statement of meristic features and other diagnostic characters, some of which are illustrated, except for the three species whose type-locality is Madeira or whose type- material includes Madeiran species. For two of these, Gohius gusfeveni and Thorogohius ephippiatus, Tables of body proportions and sensory papillae counts are provided, based on local examples, while a full account of G. maderensis follows this section. Terminology and methods of counting and measuring are as adopted by Miller ( l973a, 1974).

Gobius niger L., I758

(?) Gobius niger L.: Drouet, 1861: 412; Hilgendorf, 1888~: 208; Jordan & Gunn, 1899: 346.

366 P. J . M I L L E R

Gohius jozo L.: Pellegrin, 1913: 116; 1914: 79; Metzelaar, 1919: 281; Chabanaud &

Bathygobizts jozo (L): Fowler, 1936: 1000, 1002; Cervignon, 1960: 97. Gobitrs (Gobius) niger L.: Albuquerque 1954; 888, fig. 374. Gobizrs niger L.: Miller, 1 9 7 3 ~ : 483 (Clofnam no. 162.1.1.); Brownell, 1978: 125; Brito

& Lozano, 1980: 17 1 , figs 2, 3; Castillo & Brito, 1982: 394. Material. CANARIES: 5 males, 60+ 16 to 74+20 mm, and 1 female, 47 + 1 1 mm, from

Playa de las Teresitas, Tenerife, 1979 and 1980, coll. A. Brito; 2 males, 37.5+9.5 and 39.5 + 10 mm, from Playa de Marfolin, c. 26 km SSW Playa Blanca, Fuerteventura, 20 August 1979, coll. J. K. Dooley.

Identification. A widely distributed, euryhaline, Mediterranean-Lusitanian and eastern Atlantic boreal species, the black goby, G. niger, has been reported from Madeira (Lowe, 1839), the Azores (Drouet, 186 1 ) and the Canaries (Jordan & Gunn, 1899). African continental records exist for Morocco (De Buen, 1930; Dollfus, 1955; Brownell, 1978), Baie du Levrier, Mauritania (Pellegrin, 191 3, 1914; Chabanaud & Monod, 1927), and off Senegal (Cervignon, 1960). However, the early identification by Lowe ( 1 839) refers to Muuligobius tnaderensis, which Lowe regarded as a variety (“P”) of G. niger, while Drouet ( I 86 1 ) and Jordan & Gunn ( 1 899) may very well have applied the name to individuals of G. paganellus in the Azores, and to either M. rnaderensis or G. paganellus from the Canaries. These are both dark-coloured species of comparable size, the former common in the Azores and both so around the Canaries (see below). Nevertheless, the occurrence of G. niger at the Canaries has recently been confirmed by Brito & Lozano (1980), who describe and illustrate examples of this species from north-eastern Tenerife. Subsequently, G. niger has been found at Las Palmas, Gran Canaria (Castillo & Brito, 1982) and Fuerteventura (J. K. Dooley, pers. comm.). The present author has subsequently examined six Tenerife specimens, kindly sent by Dr Brito, and has confirmed that these show all the characteristic features of G. n i p r from European localities. Thus, fin-ray and lateral scale counts of D2 1/12, A 1/1 I (10-12), P I8 (1 7-20), and LL 38-39 ( 3 3 4 l), fall within an overall range of D2 1/12-13 (1 1-1 3), A I l l 1-12 (10-13), P 17-19 (15-20)and LL35-41 (32-42)(Miller& El-Tawil, 1974; Miller, In press). Vertebral number (including urostyle) is 28, the usual value for Gohius species. The lateral-line system (Fig. 2) displays an unbroken suborbital row d, row .Y’ ends behind the vertical of pore P . pore a lies behind the eye on the course of the oculoscapular canal, and the head canal pores tend to be wide. Head squamation includes a scaled nape but no cheek scales. In other respects (Fig. 3), the anterior nostril possesses a more or less acute triangular lappet from the rear rim, pectoral free rays are moderately developed (involving the four uppermost elements), and the pelvic anterior membrane is without prominent lateral lobes. In the male examples, the first dorsal rays are elongate, extending back, when depressed, to at least the middle of the second dorsal base. In coloration (Brito & Lozano, 1980: fig. 3), dark blotches are present in the upper anterior corner of each dorsal fin, except in the second dorsal fin of the smallest (female) example, and in the male the dark lower part of the branchiostegal membrane contrasts with adjacent paler areas of the breast and head. Two further specimens, from north-west Fuerteventura, have also been made available to the author by Dr J. K. Dooley; these juvenile males resemble the Tenerife examples, and have D2 111 1 and 1/12, A I / I 1 , P 18 and LL 40.

These definite identifications of G. niger from the Canaries would also suggest that records of this species from the adjacent shelf waters of Morocco may be accepted, but those from

Monod, 1927: 284; Dollfus, 1955: 71; Postel, 1959: 164.

CiOBllD F I S H E S OF- T E M P E R A T E M A C ' A R O N E S I A 367

2 mm -

FIG. 2 . Gohiics lii,ycr. Lateral-line sensory papillae and canal-pores (cross-hatched) of head in male, 74.0 + 20.0 mm. Tenerife: (a) dorsal and (b) lateral view. A N . PN, anterior and posterior nostrils; nomenclature as text.

368 P. J . MILLER

I m m

2 mm

AM ( d )

FIG. 3. Gohius niger. (a) Pectoral fin upper edge with free rays, in relation to first dorsal fin, in male, 72.0 + 19.0 mm. Tenerife; (b) and (c), snouts of(a) and male of Fig. 2, respectively, in oblique dorsal view from rear. AN, PN, anterior and posterior nostrils; E, eye; UL, upper lip; (d) pelvic disc in (a); AM, anterior membrane.

U

farther south still deserve confirmation against the tropical West African Gohius senegumbiensis Metzelaar, 1919, another species with long first dorsal fin rays (Poll, 1959; fig. 54) and known to be present off the Mauritanian coast (Miller, In press). Re-examination of the specimens listed by Poll (1959) shows that the pattern of head sensory papillae and canal pores in G. senegumhiensis also closely resembles that of G. niger, but that meristic values of D2 1/14, A 1/13, and LL 47-51 in the former species should clearly distinguish this from G. niger in any area of distributional overlap.

GOBlID FISHES O F T E M P E R A T E MACARONESIA 369

Habitat. On European Atlantic and Mediterranean coasts, G. nigw is frequently associated with brackish water (Miller, 1973a), but can also tolerate hypersaline as well as normal marine conditions (Botros, 197 1). Occurrence around the Macaronesian islands under marine salinities may be extensive. The known habitats are dissimilar. On Tenerife, the species was found in Cymodocea beds on muddy sand at 2-3 m (Brito & Lozano, 1980), the Fuerteventura specimens were obtained from a deep intertidal pool (J. K. Dooley, pers. comm.), and at Las Palmas, Gran Canaria, G. niger has been recorded in a settlement tank of a desalination works, again on muddy sand at mostly 2-3 m (Castillo & Brito, 1982).

Cobius auratus Risso 1810

(?) Gobius auralus: Roule & Angel, 1933: 84; Zugmayer, 1933: 84; Fowler, 1936: I3 18;

Gobius uurati4.s: Castillo & Brito, 1983: 39 I , figs 2, 4. Material. CANARIES: see Castillo & Brito (1982): none available to present author. Identi/zcation. A species of Gobizrs recently collected from a settlement tank at a desalina-

tion plant near Las Palmas, Gran Canaria, has been identified as G. airrutus by Castillo & Brito ( 1 982). The distinguishing features of this Mediterranean-Atlantic species, and others associated in a "G. auratus complex", were discussed by Miller & El Tawil (1974). The true G. aiiratus is believed to be characterized by the possession of a divided suborbital row d. LL 44-50, P 18-20, A 1/14, anterior nostril with a triangular lappet on posterior rim, a deeply emarginate pelvic disc with vestigial anterior membrane, a pair of dark cheek spots, a deeper than long, pale edged, dark mark on the upper pectoral base, and five dark blotches more or less in evidence along the lateral midline below the second dorsal fin (Miller & El Tawil, 1974). The species is known from the Mediterranean (Miller, 1973a) and unconfirmed records exist for the Atlantic coast of north-west Spain (Albuquerque, 1954; Lozano Y Rey, 1960; Iglesias, 1981) as well as from Gran Deserta, Madeira (Roule & Angel, 1933; Zugmayer, 1933). The latter may just as easily apply to G. gasteveni Miller as to the true G. auratus.

Using details given by Castillo & Brito (1982), the gobies from Las Palmas have a divided suborbital row d, longitudinal body markings and a squarish pectoral spot, a combination suggesting either G. bucchichi Steindachner or G. jullax Sarato. However, meristic data, especially P 15-16 and LL 40-43, are more reminiscent of G. couchi from the western English Channel, since Mediterranean G. bucchichii has LL values of more than 50, and G. ,fullax P 18-1 9, but G. couchi does not possess longitudinal series of small dark spots along the body. Three features shown by Mediterranean G. auratus-a deeply cleft pelvic disc, a golden coloration, and P 18-30-are not reported by Castillo & Brito (1982). It is therefore desirable that further study be made before the identification as G. auratus be confirmed. There is always the possibility that the population represents an undescribed species, but some of the disparity with better known Mediterranean forms may be due to meristic trends like those shown by G. paganellus, with low LL counts seen in Canaries examples of the latter (Fig. 8). The same trend applied to G. bucchichii could approach the LL count of 40-43 found by Castillo & Brito ( 1 982), although Brownell (1978) notes 48-5 1 scales in LL for Moroccan examples of G. hucchichii.

Albuqerque, 1954: 890.

3 70 P. J . MILLER

C

Fic. 4. Gobius gasteveni. Lateral-line sensory papillae and canal-pores (cross-hatched) of head in male, 54.0 + 15.0 mm (MMF 13096A), Funchal, Madeira; (a) dorsal, (b) lateral view. Abbreviations as Fig. 2.

GOBIID FISHES OF T E M P E R A T E M A C A R O N E S I A 37 1

Gobius gasteveni Miller, 1974

(?) Gobius auratus Risso: Roule & Angel, 1933: 84; Zugmayer, 1933: 84; Fowler, 1936:

Gobiirs gasteveni Miller, 1974: 468, fig. 1, pl. 1 (England: off Plymouth, Devon); Tortonese 13 18; Albuqerque, 1954: 890.

& Hureau, 1979: 37 (Clofnam no. 162.1.7a).

T A B L E I Bodj. pruporiion 01' Macaronrsian Gobius gasteveni and Thorogobius ephippiatus. For meihod o f mcwsiiring. see Miller 11969J: individual values given ,/or C. gasteveni; ihosr 1i)r Th. ephippiatus are range and. in

parc~nihesis. rnean i S.D.

Species

Sex Male Male Males SL (mm) 54.0 42.5 75.0-1 02.0 n I ' 1 2 43

Vo SL, H Hw SN/DI SN/D2 SN/AN SN/A SN/V C P Dlb D2b Ab CI PI VI Vd Ad Aw C Pd V/AN

Okr CP. CPd

Yo H, SN E PO C H d Hw

Yo E, I

Yo V/AN, V

25.0 14.8 31.5 51.9 50.9 55.6 27.8 19.4 12.0 29.6 26.9 27.8 24. I 19.6 17.0 17.2 14.8 11.7 20.7

60.0

20.7 32.6 51.9 23.0 59.3

18.2

94.6

27.5 14.6 32.9 52.9 51.8 56.5 29.4 18.8 11.8 31.1 25.9 25.9 26.4 20.7 17.7 16.9 14.8 10.4 21.7

55.0

19.7 34.2 47.0 23.9 53.0

17.5

95.7

26.5-26.7 14.2-1 5.5 34.3-3 5.9 52.9-54.4 5 3.9-54.7 57.8-60.9 284-30.3 22.8-23.9 11.3-12.0 244-26.7 16.7-20.0 26.0-28.7

224-24. I 245-2 7.7

16.9- 19.4 15.7-16.4 9.C-14.4

10.2-1 2 . I 24.1-25.5

43.5-50.5

22.5-32.7 304-33.9 44.8-50.0 2 8.6-32.2 5 3.7-58.4

12.7-33.3

88'5-1 00.0

(26.6-10.08) ( 14.9 k 0.46) (34.8 k0 .64) (53.5k0.56) (54.4-10.29) (59.1 k 1.13) (29.0-10.91) (23.4 k0 .41) ( I 1 .7k0.27) (25.5 10 .96 ) ( 1 8 . 4 i l . 1 9 ) (27. I k 1.02)

(23.0 -1 0.80) (18.3-10.92) (16.1 10 .27 ) ( I 2.4 k 2,05) ( I 0.8 I 0 . 7 6 ) (2 5.0 k 0.5 8)

(46.1 5 2.64)

(26. I -14.05) (32.4 k I ,34) (47 .9k 1.91) (30.5k 1.48) (56.0k 1.67)

(2 1 .O k 8.89)

(92.2 k4 .67)

(25.7 _+ I . 22 )

'MMF l3096A: 'MMF22975; JMMF 181, 16959, 16960,21491

372 P. J . MILLER

TABLE I I Numbers ofsensory papillae in head laieral-line syslem of Macaronesian Gobius gasteveni and

Thorogobius ephippiatus

Thorogobitts Thorogobiu \ Species Gobius gaasteveni ephrppiarus Species Gobius guarievrwi ephipprutiic

Sex Male Male Males Sex Male Male Males

n 1 1 1 2 43 n 1 ' 1 2 43 SL 54.0 42.5 75&102.0 SL 54.0 42.5 754-102.0

PO: rl r2 S I .S2

s3

c2 c1 c2 CI

so: I 2 3 4 5 6 7 b d

P M : e i f

3,4 3 ,3 3,3 3 ,3 4,2 3 ,3 4 ,4 3 ,3 5,4 3 , 3 9 .9 6 , 4 6 ,5 6 ,6

12.9 8 ,8 5,4 4,3

14, I I 11, 10 8 , l O 7.7 8 ,9 9 ,6 8.8 8.6

1 0 , l O 10,lO 12, 10 13, 12

ab ab 14,15 10, I I 29,29 24,21

65,62 52,51 29,27 30,30

7,4 5,6

8-1 I 10-13 4-13

11-16 5-8

17-25 8-1 I

11-15 5-6

11-29 I 7-2 I 15-22 I 7-25 20-25 38-45 ab-2

3 5-40

14-95 3 4 4 3 2 5-2 7

26-3 1

0s: x' X2

4 ir I' as1 as2 as3

la I

la2 la '

O P 01

oi

AD: n g

m h

Z

0s

0

14,14 10.13 a ,4 8 , 5 6 ,6 5 , 5 2,2 4 ,4 ab ab a, I 3 ,2 a.6 7 ,6 a, 8 8 , 9 a .8 10, 10 a, 2 3 ,3 a.2 2 ,2 ab ab

31,30 27,30 I I , 14 12, I3 7 ,7 8 , l

5, 5 6 ,6 8.7 8 ,7 3, 5 6 ,7 5, 5 6 ,4 a ,9 10, 1 I

2 1-26 8-10

12-16 6-9 1-2 1-2

16-19 c 13

22-26 4-6 8-9 ab

35-45 16-24 10-12

13-19 19-23 6-9( 20) 8-10

25-1 I

1, 2 , 3, As Table I; a, abraded; ab, absent.

Material. MADEIRA: 2 males, 42-5+ 1 1 and 54+ 15 mm (MMF 22975, 13096A) from Funchal, 8 November 1971 and 29 November 1957.

CANARIES: one male, 45.5 + 1 1.5 mm, and one female, 34.5 + 8.7 mm (ZMUC), 3.5 miles S, 0.5 miles E Las Palmas, Gran Canaria, 25 March 1930, Th. Mortensen; one male, 44.0 + d mm (ZMUC), off La Luz, Gran Canaria, 2 April 1930, Th. Mortensen.

Identification. This typically offshore species was described by the author (Miller, 1974) from material collected chiefly in the western English Channel, but also including two specimens from Madeira, recognized by Mr G. E. Maul at the time of capture as belonging to an unusual form. An earlier Madeiran record of a goby, taken by line at Deserta Grande and identified as G. uurutus Risso (Roule & Angel, 1933), may denote this species; comparison between G. gusteveni and other close relatives, including G. uurutus, was made by Miller

GOBIID F I S H E S OF T E M P E R A T E M A C A R O N E S I A 313

& El-Tawil (1974). As noted above, the new record of G. uurutus, from Gran Canaria (Castillo & Brito, 1982) requires confirmation but does not appear to be G. gusteveni, because of differences in body coloration, shape of pectoral spot, and pectoral counts of merely 15-16. Examples of true G. gusteveni are now reported from the Canaries.

A full systematic description of G gusteveni was given by Miller (1974), but body proportions and sensory papillae numbers in the Madeira examples are here supplied in Tables I and 11. Diagnostic features for this species comprise divided suborbital row d (Fig. 4), LL 4 0 4 5 (37-49, anterior nostril with single thin process (Fig. 5 ) , pelvic disc rounded to emarginate posteriorly (Fig. 5) , white cheek mottling, a longer than deep mark on the upper pectoral base, and four lateral blotches below the second dorsal fin (Miller, 1974: pl. I). Other meristic characters, from all material of this species, are D1 VI (V-VI); D2 1/14 (13-15);

2 m m U

( c )

I

AM

FIG. 5. Gobiic.~ gastcwnr (a) snout of male of Fig. 4, in oblique dorsal view from rear; (b) pectoral fin upper edge with free rays, in relation to first dorsal fin in same fish; (c) pelvic disc in male, 42.5 + 11.0 mm ( M M F 22975); abbreviations as Fig. 3.

374 P . J. M I L L E R

A 1/13 (12-14); P 19-21 (18-22); PD 23-28 (21-29), and VC 28 (including urostyle); the Madeiran and Canaries examples show D1 V1, D2 1/14, A 1/13, P 19-20, LL 41-43, PD 20-22 and, from radiography, VC28. These individuals show greater freeing of uppermost pectoral rays than in northern specimens (Miller, 1974, fig. 2; Fig. 5) . In life, Mr G. E. Maul noted that one of the Madeiran fishes (Miller, 1974, pl. I) was pale grey above, and white below, with reddish brown lateral blotches, and, on the cheek and opercle, had white spots, set in orange on the latter. There were no dark bands across the dorsal fins but the anal fin and lower caudal fin had a narrow dark grey edging. Otherwise, the dorsal fins and the upper two-thirds of the caudal fin had reddish-brown rays, while the pectoral spot was mottled dark and reddish brown and the upper lobe of the pectoral fin orange.

Habitat. In the western English Channel, G. gasteveni has been trawled at depths from 36-74 m, usually on muddy sand but also on shell or small stones. At Madeira, one specimen (MMF 13096A) was taken in Funchal Harbour in 35 m and the second (MMF 22975) caught in a nylon pot at about 100 m depth in Funchal Bay. The specimen identified as G. auratus by Rode & Angel (1933) was collected at Deserta Grande in 2-12 m. Off Gran Canaria, the Las Palmas examples were dredged in 110-270 m, on hard ground, while the male from La Luz was dredged in about 180 m. So far, G. gasteveni is known only from areas at opposite ends of the eastern Atlantic Lusitanian-Mediterranean (Ekman, 1953) or Lusitanian (Briggs, 1974) faunistic province, but there is little doubt that further collecting will disclose its presence throughout the intervening shelf waters of Morocco, Iberia and the Bay of Biscay.

Gobius paganellus L., 1 758

(?) Gobius niger L.: Drouet, 186 1 : 94; Hilgendorf, 1888a: 208; Jordan & Gunn, 1899: 346. Bathygobius paganellus (L.): Fowler, 1936: 1000,1003, I3 16 (part); Noronha & Sarmento,

1948: 154; Nunes, 1953: 77, 78; Collins, 1954: 130. Gobius paganellus: Giinther, 186 1: 52; Steindachner, 1868: 4 13 (part); Hilgendorf, 1888,:

208; 18886: 79; Collett, 1890: 222; Vinciguerra, 1893: 32 1 ; Boulenger, 19 16: 35 1 ; Metzelaar, 1919: 381; Brownell, 1978: 124; Arruda, 1979: 18, figs 5-7: 1980a: 340, 345, 346; Brito & Lozano, 1980: 175; Castillo & Brito, 1982: 394.

(?) Gobius rnaderensis Val.: Regan, 1903: 348. Gobius (Macrogobius) paganellus: Albuquerque, 1954: 890. Material. AZORES: 5 males, 65+ 145-100+23.5 mm, and 5 females, 87.5+ 19-

95 + 2 1.5 mm, 1972, coll. University College, London, expedition (BMNH, uncatalogued); 2 males, 39 + 9 and 45 + 10.5 mm, and 2 females, 37.5 + 8 and 45 +9 mm, from Populo, Siio Miguel, August 1972, coll. UCL expedition; 5 of indeterminate sex, 25.5 +6.0- 33.5 + 8.0 mm, from Fayal, August 1978, coll. L. Arruda; 2 males, 69 + 16 and 93 + d mm, and 3 females, 60 + 13.5-85 + 22 mm, from Faja de Caldeira, S2o Jorge (38'38 'N, 27'56 ' W), 13 June 1981, CANCAP-RMNH; one juvenile, 21 +5.5 mm, from Santa Cruz, Flores (39'27 'N, 3 1'07 'W), 9 June 198 1, CANCAP-RMNH; one male, 92 + 2 1 mm, from Caloura/ Vila Franco, SBo Miguel(37O43 'N, 25'20'W), 19 September 1979, CANCAP-RMNH; one male, 72 + 12.5 mm, from Baia da Praia, Santa Maria (36'57 'N, 25'06 'W), I5 September 1979, CANCAP-RMNH; one male, 59 + 13.5 mm, from Porto Pim, Horta, Fayal(38'3 1 'N, 28'38'W), 8 October 1979, CANCAP-RMNH; one male, 59+14mm, from Baia San Lourenco, Santa Maria (37'00'N, 25'03 'W), 16 September 1979, CANCAP-RMNH; one

GOBllD FISHES OF T E M P E R A T E M A C A R O N E S I A 315

male, 70.5 + 18 mm and 7 juveniles, 18.5 +4.5-22 + 5 . 5 mm, from I. da Villa, Santa Maria, 29 May I98 1 , CANCAP-RMNH.

MADEIRA: one male, 57+ 14 mm (BMNH 53.9.6.1), “purchased of S. Steven’s’’; 5 males, 51 + 13-83+ 19 mm and 2 females, 84+22 and 85+21 mm (BMNH 53.12.26.5 & 7, 58.4.4. 15-17, 131); one, sex indeterminate, 73.5+ 17 mm (BMNH 58.6-12.4) [from] Johnstone; 3 females, 64+ 17.5-72+ 18 mm (BMNH 99.1.16. 11-16), [from] A. J. Arundell; one male, 48.5+ 12.0 mm (BMNH 1936.1 1.28.7), [from] Carter; 39, sex indeterminate, 23.0+ 6.0-50.0+ 13.5 mm (BMNH 1953.1 1.1.300-387 (part)), from Baia do Salto do Cavale, Funchal, 24 August 1953, coll. D. W. Tucker; 16 males, 30.5+7.0- 78.5+ 19-0 mm, 6 females, 40.0+ 104-44.5+ 10-5 mm; 7 sex indeterminate, 18+4-43.5+ 10.5 mm, from Ponta Gorda, c. 2 km W. Funchal, 25 July 1975 (A. C. Wheeler); one male, 36.0+8.5 mm, Barreirinha, c. 0.5 km W. Funchal, 26 July 1975 (A. C. Wheeler); one male, 24*5+6.0 mm, one female, 37.0+9.0 mm, and one, sex indeterminate, 32.0+7.5 mm, from Pta does Reis Magos, Canico, c. 8.5 km E. Funchal Harbour, 30 July 1975 (A. C. Wheeler).

CANARIES: 9 males, 37.5+ 10.5-70.0+ 17.0 mm, and 7 females, 36*0+ 104-68.0+ 16.5 mm (BMNH uncat.), A. K. Totton; one, sex indeterminate, 25.0+d mm (BMNH 1952.4.3.1 .), Arrecife, Lanzarote, C. Mallabey; 4 males, 54+ 12-60+ 13.5 mm, and 2 females, 48 + 1 1 and 53 + 12 mm (MHNG 1 561.66-7 I ), from Playa del Ingles, Maspaloma, Gran Canaria, 24 December 1975, coll. P. Haymoz; 3 males, 47+ 11-62+ 12.5 mm, and 3 females, 4 1 + 9-78 + 18.5 mm (MHNG 1572. 5 1-100, 1573.1-3), from Playa de Los Cristianos, Los Cristianos, Tenerife, 1 1 January 1978, coll. P. Haymoz; one male, 32.5+7.5 and one female, 37.5+9 mm from Los Cristianos, 28 May 1979, coll. M. Linley; one male, 52 + 13.5 mm, Tenerife, April 1980, coll. M. Linley.

Ident$cation. The rock goby, Gobius paganellus, is a warm-temperate species, widespread throughout the Mediterranean, as well as in the Black Sea, and along the eastern Atlantic seaboard, from western Scotland to Senegal, with a doubtful record from Pointe Noire, Congo Republic (Miller, In press; Roux, 1957). In the Macaronesian islands, G. puganeflus was first recorded by Giinther ( 1 861 ) from Madeira, and, for the Canaries, at Tenerife, by Steindachner (18681, although much of the latter’s account of G. paganellus there and at Madeira obviously refers to Mauliguhius maderensis, regarded as a variety of the former species. This misidentification was quoted by Vinciguerra ( 1 893), and the same mistake was made by Collett (1 890) whose Funchal paganellus would seem to be nzaderensis. However, Boulenger (1916) notes further examples of G. paganellus from Madeira as well as Great Salvage, to the north of the Canaries. Abundant material, listed above from Madeira and the Canaries, shows that the rock goby must be of frequent occurrence around these islands.

For the Azores, a record of G. niger from San Miguel by Drouet ( 1 86 1 ) probably relates to G. paganellus. The species was also listed by Hilgendorf ( I888a, b) whose brief description agrees with recent examples from these islands. Later records are provided by Regan (l903), whose “G. maderensis” is very likely G. paganellus, and by Fowler ( 1 936). Modern work by Arruda (1979) has shown that this goby is numerous on the south and eastern coasts of Fayal, with CANCAP specimens (see above) from other islands (Santa Maria, Siio Jorge, Sad Miguel, and Flores).

Intertidal and sublittoral on more sheltered rocky shores, G. pagunellus, with D2 1/13-14, A 111 1-12, P 21-22 (20-23), is characterized by LL normally 50-56 (48-57), well-developed free pectoral rays, rounded pelvic disc with more or less distinct small lateral lobes on the

,

AM

GOBl lD FISHES O F TEMPERATE MACARONESIA 371

FIG. 7. Gohiics pugu:rrnc//irs. Lateral-line sensor! papillae and canal-pores (cross-hatched) of head in male. 78.5 + 19.0 mm. Madeira. Abbreviations as Fig. 2.

anterior membrane, anterior nostril with digitate process from the rear margin (Fig. 6) and, in the lateral-line system (Fig. 7), pore a at the end of a short suborbital extension from the oculoscapular canal, a divided suborbital row d, and row .vl ending behind pore j?. Coloration is usually a dark mottling over fawn, becoming more uniformly dark brown or purplish in breeding males, but always including a pale to orange horizontal band across the upper edge of the first dorsal fin (Miller, In press).

The general features of most Macaronesian G. pagarzellus agree well with the above diagnosis, based on Mediterranean and north-eastern Atlantic material. External characters (Fig. 6) and the head lateral-line system (Fig. 7) are illustrated for specimens from Madeira. In the male collected at Tenerife in 1980, life colour of the superior pale band across the first dorsal fin was noticeably more reddish than is usually seen in fish from north-western Europe. Material from the Azores includes several examples with the first dorsal pale band much narrower than in European populations. Also, counting fin-rays and scales in the Macaronesian G. paganellits revealed many values below 50 for lateral scale series in Canary

FIG. 6 . Go/Jil~,s pu,gunc~//irs (a-c) snouts of (a) female. 78.0+ 18.0 m m (MG 1572. 51-100. 1573. 1-3. part). Tenerife, (b) female, 44.5 + 10.5 mm. Madeira. and (c) female. 43,0+ 10.5 mm. Madeira, in oblique dorsal view from rear: (d) and (e). pelvic disc in juvenile. 25.0+6.0mn1 (BMNH lY53.1 1.1.300-387. part). and male. 78.5+ 19.0 mm, respectively, both from Madeira: abbreviations as Fig. 3.

378 P. J . M I L L E R

D2 12 13 14 15 I I .k S.D. 2.24 S.E.

C I 24 2 - 27 13.04 0.33 0.14 M 3 43 I - 47 12.96 0.29 0.09 A - 6 24 - 30 13.80 0.40 0.16 Me I 26 35 - 62 13.55 0.53 0.15 Ch - 18 71 I 90 13.81 0.42 0.10

A 10 I 1 12 I3 I I 4 S.D. 2.24 S.D.

C - 2 6 - - 26 12.00 - - M - 42 2 - 44 11.05 0.21 0.07 A - 2 27 I 30 11.97 0.32 0.13 Me I 19 41 - 61 11.66 0.51 0.15 Ch - 26 63 - 89 11.71 0.46 0.11

P 18 19 20 21 22 23 n .k S.1). 2.24 s.1:.

c - - 2 16 29 5 52 21.71 0.69 0.21 M - - I 7 32 7 47 21.96 0.62 0.20 A _ _ _ 10 30 18 58 22.14 0.68 0.20 Me - - 2 29 52 19 102 21.86 0.73 0.16 Ch 2 30 41 49 16 - 138 20.34 0.99 0.19

.k S.D. 2.24 S.E. LL 46 47 48 49 50 51 52 53 54 55 56 57 58 59 n

C 2 3 6 5 7 1 1 1 0 3 I - - - - - 48 50.21 1.94 0.53 M - 2 - I 2 4 8 6 1 0 4 2 - - - 39 52.64 2.08 0.75 A _ _ - - I I 2 4 13 14 8 4 6 3 56 55.18 1.95 0.58 Me - - I 2 8 1 1 2 2 2 2 3 6 1 8 9 1 I - 131 53.21 1.79 0.35 Ch - - I 4 4 6 19 26 37 12 8 4 - - 121 53.31 1.77 0.36

specimens, and a low modal value in second dorsal and anal rays for both Canary and Madeiran fishes. These discrepancies between Macaronesian data and the usual distribution of meristic values for European populations suggested the need for more detailed comparison with samples available from the coasts of the western Mediterranean basin and from the western English Channel. Material and results are shown in Table I11 and Fig. 8.

All five populations overlap in range of values for second dorsal, anal, and pectoral fin- rays, and number of scales in lateral series, so that, by these criteria, they can be regarded as belonging to the same morphological species. However, analysis of variance indicates some significant differences between mean values for the characters used, although it should be noted that second dorsal and anal counts show a high degree of correlation. Canarian and Madeiran G. paganellus have significantly lower means for second dorsal and anal counts

GOBllD FISHES O F TEMPERATE MACARONESIA 379

and the former also differ significantly in the low mean value for scales in lateral series. Among the populations, the Azores material has the highest means for anal, pectoral and scale counts, separated in those for median fin and lateral scale counts from the Canarian and Madeiran specimens, but shows some relationship in median fin ray values with those from the Mediterranean and Western Channel. Madeiran G. pagunellits do not differ from western Mediterranean examples in means of pectoral and scale counts, while the Canaries population retains only similarity in pectoral ray values. Brownell ( 1 978) has given second

Me

I

n

D2 14 ‘‘1 15

13

C M A Me C h 27 47 30 T T

26 44 30

1

62 90

T

n 5 2 47 58 102 138

19 0.0. 0 00. m u 0.

22 000 . 23 .... 24

n 48 39 56 131 121 4 5 r 7

60

FIG. 8. G‘ohIii.s pagnnc~//zi.s. Variation in number of second dorsal branched rays (D2). anal branched rays (A). pectoral rays (P). and scales in lateral series (LL). between five areas (material as text or indicated): A , Azores, C, Canary Islands; Ch, Western Channel ( I . Helford (32 specimens); 2. Falmouth (6); 3. Looe (23); 4. Wembury (38)); M , Madeira; Me, Western Mediterranean ( I , Alicante (7): 2. Banyuls (7): 3, Villefranche-sur-Mer ( 5 ) ; 4, Portofino (27); 5 , Sestri Levante (2); 6. Giglio (19). Graphic comparison (‘Dice-grams’) based on Eberhardt (1968); for each sample is given range (vertical line), mean (long horizontal line). and, on each side of mean, one standard deviation (open rectangle) and 2.24 standard errors (black rectangle); n, number of observations. Matrix for each character indicates significant (black squares) or non-significant (open squares) diferences between localities at 0.05 level in GT2 method for multiple comparisons among pairs of means (Sokal & Rohlf, 198 I ) .

380 P. J. M I L L E R

dorsal, anal and pectoral fin-ray counts for G. paganellus collected on the Moroccan coast between Agadir and Rabat. These values (D2: X 13.9; S.D. 0.36; A: X 11.92, S.D. 0.44; P: X 2 1.24, S.D. 0.71) indicate close resemblance to the Western Channel and Azores populations in median fin-ray elements, but a significantly distinct mean for pectoral rays between that of the Canaries and the Western Channel. These findings are further discussed in a later section.

Arruda (1979), as well as providing a table of body proportions for Azores G. paganellus, has also compared their meristic ranges with values given by previous authors (Arruda, 1979: table VI). Ranges indicated by Arruda for the Azores material agree with present findings, but the usefulness of such data from the literature is limited without the opportunity to stan- dardize results from different counting methods, and the provision of further basic statistical information such as, at least, frequency distribution of values. As a practical criterion for species determination, it should be noted that the minimum count of 46 scales in lateral series for Canaries G. paganellus is still well above the values of 39 and 40 found in a small relative, G. ater Bellotti, known, as yet, only from the Mediterranean (Miller, In press).

Arruda (1979) has also commented on difference in suborbital papillae pattern between the Azores G. pagunellus and examples from elsewhere, as depicted by Goren & Klausewitz ( 1 978) from the Mediterranean and Red Sea (immigrant). The Madeiran specimen of this species illustrated here (Fig. 7 ) has a juxtaposition of upper and lower parts of rows 5 and 6 more like that shown in Goren & Klausewitz’s drawing than the illustration by Arruda of the pattern in a rock goby from the Azores. However, small examples from the latter islands, sent to the author by Dr Arruda, have a configuration of rows 5 and 6 similar to Mediterranean specimens. In this particular feature, variation between individuals from the same locality may be encountered. It should also be noted that the Red Sea specimen shown by Goren & Klausewitz (1978) is very unusual-unique in the author’s experience of continental G. paganellus-in displaying a continuous row d. Another specimen (HUJ 4699), from the same place, has the normal gap in the horizontal section of this row. By coincidence, a few CANCAP G. paganellus from the Azores have been found to have the usual gap in row d very narrow or entirely occluded, although others display the typical subdivision.

Habitat. Arruda (1979) has shown that, at Fayal in the Azores, G. paganelhis is common in the lower intertidal zone of shores with abundant weed, or in pools beneath overhangs and boulders on more exposed shores, the young fishes being of sporadic occurrence at higher levels. The intertidal fish community at Fayal has been compared with two on the Portuguese coast by Arruda (19804. At Madeiran localities, G. paganclltrs occurs in shore pools with M . maderensis (see below), while Brito & Lozano (1980) note that, at Tenerife, G. paganellus inhabits sublittoral rocky ground, as well as the intertidal zone (to which M. maderensis is restricted).

In the field, difficulty may be experienced in distinguishing small G. paganellus from M. maderensis, of comparable size and collected from the same shore pool. Diagnostic features of coloration and head lateral-line system are summarized in Appendix I . The precise ecological relationships of these two sympatric gobiid species, with probably similar life- history, would merit further study.

Lesuenrigobius suerii (Risso, 181 0) Gohizrs Iesueurii Risso: Vaillant, 1888: 18, 348. Lesucurigohiu.s sirerii (Risso): Miller, 1 9 7 3 ~ : 499 (Clofnam no. 162.16. I ) .

GOBIID FISHES OF T E M P E R A T E MACARONESIA 38 I

Material. CANARIES: see Vaillant (1888); none available to present author. Ldent$cation. The eastern Atlantic-Mediterranean genus Le,sireirrigobiir.s Whitley, with

longitudinal suborbital and interorbital papillae rows a, ( a , and p , no head canals, low number of scales in lateral series, compressed body, and lanceolate caudal fin, comprises five species. inhabiting coastal and offshore shelf waters and associated with finer deposits, in which burrowing has been described for L. . J k s i i by Rice & Johnstone ( 1 972). Lesireurigohiiis suerii, the type-species, is widespread in the Mediterranean, including the Alboran Sea, and also known from the Atlantic coast of Morocco (Miller, 1 9 7 3 ~ ; Vaillant, 1888, Dollfus, 1955; Peres, 1964). In contrast to the other two Mediterranean-Atlantic forms, L. sanioi (De Buen, 19 18) and L.,/rie.sii, L. szicrii lacks scales on the nape and displays oblique yellow and blue bands across the head and body. Meristic formula is DI VI, D2 1/13-14, A 1/13-14, P 17-20, LL 26-28, and VC 27 (Miller, In press).

To date, the only evidence for the occurrence of this goby in Macaronesian waters is supplied by Vaillant (1888), who lists a specimen obtained off the Canaries by the "Travailleur" ( I 882, stn 47 at 28'28 '30 "N, 18'32 'W). No recent material is to hand, and there is a possibility of confusion with the following species or L. s u n x i , known in the Atlantic from Portugal (Almeida & Gomes, 1978) to Mauretania (De Groot & Nijssen, 197 I , as Gobius pagunellus).

Habitat. The bathymetric range of L. .sirerii extends from inshore to 220 m, on mud or muddy sand (De Buen, 1923). The Canaries specimen noted by Vaillant (1888) was caught in 80 m. on sand and rocks.

Lesuenrigobius heterofusciutus hdaul, I971

Lesueurigohius hetrryfasciatits Maul, 197 I , fig. 2 (Morocco: W. of Agadir, 3 1'35 'N, IO"10.5'W); Miller, 1 9 7 3 ~ : 500 (Clofnam no. 162.16.3); Maul, 1976: 48, fig. 22; Tortonese & Hureau, 1979: 38.

Material. MADEIRA: see Maul ( 197 1 ). N o further examples available to present author. Ldentjfication. The present species of LesireiiriKohizis was fully described by Maul ( I 97 1)

on the basis of two examples from Morocco (SL 38.5 and 42 mm) and one from Madeira (SL 44 mm). It resembles L. suerii in having no predorsal scales, and separate anterior dorsal papillae rows g and h, but may be distinguished by the presence of several narrow, well demarcated, dark brown vertical bands down the sides of the body, separation of suborbital row b from the preopercular groove, and more gill rakers (3+ 11-12 on the first arch) than in other species (2+8) (Maul, 1971: fig. I ) . Meristic formula is D1 VI, D2 1/14-15, A 1/14-15, P22-24, and LL28 (Maul, 1971).

Habitat. Off the Moroccan coast, L. hrteryfusciatirs was taken during the "Meteor" expedition by Agassiz trawl in 150-160 m and 345-1 70 m , while the Madeiran specimen was found inside the mouth of an Auk)pits.filamento.sit.s in Funchal fish market (Maul, 1971, 1976).

Mauligobius muderensis ( I'alenciennes, 183 71

Synonymy and material. See later section. Ldent$cation. First described from Madeira, by Valenciennes (1837), the validity of

Gobius maderensis was soon questioned by Lowe ( I 839) who came to regard the species

382 P. J. M I L L E R

as a mere variety of the European G. niger. Later, Steindachner (1868), from material obtained at Santa Cruz, Tenerife, concluded that G. rnaderensis was a local form of G. paganellus, a view repeated by Fowler (1936), and presumably leading to the omission of this goby from a list of Madeiran fish prepared by Maul (1949), and also from the work of Albuquerque (1954). Other, earlier, authors have regarded this species as distinct. Gunther ( 1 86 1 ) and Von Martens ( 1 876) noted examples from Madeira, while Vinciguerra ( 1 893) and Zugmayer (19 1 I ) recorded the species from the Canaries and the Salvage Islands, respectively. On the basis of further material from the Canaries, Brito & Lozano (1980) have recently discussed its nomenclature. A record by Regan (1903) from the Azores remains unconfirmed by later collecting (Arruda, 1979, 1 9 8 0 ~ ; CANCAP project).

Re-examination of the syntypes of G. rnaderensis, together with study of many specimens from Madeira, the Canaries and the Salvage Islands, has shown that this species is easily distinguished from other gobies, and possesses a number of morphological features, especially a well-developed suborbital row 7, which have supported the recognition of a separate genus to contain G. rnaderensis, as well as a tropical West African form, G. nigri Gunther. For this genus-group taxon, the name Madigobius has already appeared within a key and check-list by Miller (198 l), satisfying ICZN criteria for publication. In the present paper, a later section provides a full generic definition of Mauligohius. designation of type-species, key to species, redescription of Gobius rnaderensis. with synonymy and all Macaronesian records, and a consideration of phyletic affinities.

Thorogobius ephippiatus (Lowe, 1839)

Gobius ephippiatus Lowe, 1839: 84 (Madeira); Fowler, 1936: 1004; Noronha & Sarmento, 1948: 154; Maul, 1949: 155.

Thorogohius ephippiatus (Lowe): Miller, 1969: 831, fig. 2A, 3A, 4A, pl. I; Miller, Rice & Johnstone, 1973: 234, fig. 3; Miller, 1973~ : 512 (Clofnam no. 162.24.1); Almeida & Gomes, 1978: 8, pl. I1 (fig. 6); Tortonese & Hureau, 1979: 39.

Material. SALVAGE ISLANDS: one male, 92 +25 mm (MMF 18 l), May 1922. MADEIRA: 2 males, 75 + 19.5 and 102 + 27 mm (MMF 16959, 16960), Funchal Bay, 7

March 1960); one male, 97.5+28 mm (MMF21491), Funchal Bay, 13 July 1965. ldmt$cation. A gobiid whose type-locality is Madeira (Lowe, 1839), and long overlooked

by European ichthyologists, Th. ephippiatus, the Leopard-spotted goby, was independently discovered more recently by aqualung divers in the western English Channel (Corbin, 1958; Miller, 1961), and subsequently found to occur at many European localities in the temperate eastern Atlantic and Mediterranean (Miller, 1969; Miller et al, 1973; Wirtz, Hoare & Hiscock, 1975; Dunne, 1976; Almeida & Gomes, 1978). Comparison of British material with specimens from Madeira and the Salvages, in conjunction with Lowe’s original description, has established their common identity with G. ephippiatus (Miller, 1969), and sight records by Brito (pers. comm.) have now extended the southern limit of known distribution to Tenerife, in the Canaries.

This species was made the type of a new genus, Thorogobius, into which the true G. rnacdepis of Kolombatovic ( 1 89 l ) , from the Mediterranean, was also placed (Miller, 1969), and to which the tropical Atlantic Gobius angolensis Norman, 1935, must be added (Miller, 1981). The most distinctive character of Th. ephippiatus is the presence of large orange blotches over an otherwise usually pale head and body, and a conspicuous dark spot on the

FIG. 9. Thorogohiir5 oph;ppiumx (a) pelvic disc: (b) snout, in oblique dorsal view from rear; (c) and (d). lateral-line sensory papillae and canal-pores (cross-hatched) of head, dorsal and lateral view respectively. all in male. 97.5+28.0 mm ( M M F 21491), Madeira. Abbreviations as Figs 2 and 3.

384 P. J . M I L L E R

first dorsal fin (Miller, 1969: pl. I) . In contrast to species of the related genus Gohius, there are no predorsal scales, the uppermost pectoral rays are contained within the fin membrane, the pelvic anterior membrane has no marked lateral lobes, and the anterior nostril is a short tube, without a dermal process from its rim (Fig. 9, (a) and (b)). The head lateral-line system (Fig. 9, (c) and (d)) has long anterior dorsal rows g and XI, with rows o meeting in dorsal midline, and suborbital row h approaches or meets vertical row z. Typically, as in the present material, row .Y' meets row n and 6 approaches row P . Examination of meristic characters and numbers of sensory papillae in certain head rows (suborbitals 1-6, h, and d, anterior dorsal .t4 and z , opercular ot, and preoperculo-mandibular e and i) has suggested regional variation, with highest average value for pectoral rays in the Macaronesian examples, which resemble Celtic Sea and Irish specimens in tending to have more head papillae at adult size than is found in a western Scottish population (Miller el al.. 1973: Dunne, 1976).

For the present Madeiran and Salvage material, body proportions are given in Table I , and sensory papilla numbers in Table 11; their meristic characters are DI VI, D2 111 1 , A 1/10, P 19, LL 36-40 (36: 2,37: 2,38: 3,40: 1) and VC 28.

This species is an extremely wary inhabitant of inshore rocky ground, found within or near rock crevices and beneath overhangs on fine mud to muddy shell gravel, or coarse shell, gravel and pebbly substrata, situated in deep gullies, in caves, at the foot of open rock faces, and even among concrete blocks or wreckage (Miller, 1969; Barber, 1973; Wirtz, Hoare & Hiscock, 1975; Dunne, 1976). Vertical range is from rare occurrence in deep MTL-LWST pools, to depths of 3 0 4 0 m in the Mediterranean, but the goby is most frequently encountered at about 6-1 5 m, beneath the sublittoral limit of the seaweed Laminaria hyperhorea in boreal waters (Miller, 1969; Miller et al., 1973). The Madeiran examples were captured on rocky ground, at 65, 100 and 120 m, deeper occurrence than noted in European observations. At Fuerteventura, a southward extension of known range, the species has been photographed in a grotto at 20 m (P. Brito, pers. comm., 13 March 198 1).

The diet and life-history of Th. ephippiatus are described by Miller (1969) and Dunne (1976).

The genus Muufigobius Miller 1981 Mauligohius Miller, 198 I , Gobiidae 6, 8, fig. 18. Type: Gobius madevensis Valenciennes I837 (by present designation). Body moderately elongate; eyes dorso-lateral, with interorbital space about one third to

over half eye-diameter; cranial roof covered by dorsal axial musculature; caudal fin rounded, slightly shorter than head; body with ctenoid scales, original and replacement, 42-57 in lateral series; head and predorsal area, breast, upper opercle and cheek with scales; uppermost and lowermost scales of caudal peduncle not enlarged; snout convex in transverse section; anterior nostril tubular, with single or digitate tentacle on posterior rim, as long as tube; posterior nostril pore-like, near orbit; pectoral girdle without dermal flaps on anterior edge; pectoral fin with upper rays partially free from membrane; pelvic disc complete, rounded, posterior edge well before anus; pelvic anterior membrane well developed, with prominent lateral lobes; mouth oblique or horizontal, jaws subequal with posterior angle not behind pupil; upper lip width uniform, somewhat narrower than lateral preorbital area; chin without transverse fold or barbels; teeth in jaws erect, caniniform, in several rows medially; posterior median teeth of upper jaw not enlarged; outer row of teeth in both jaws enlarged, lower jaw with lateral group of larger canines present or absent; pharyngeal teeth

385

FIG. 10. Muuligobius maderensis. Lateral-line sensory papillae and canal-pores (cross-hatched) of head in female, 70.0+ 17.0 mm, Madeira. Abbreviations and nomenclature as Fig. 2.

386 P. J . MILLER

FIG. I I . Skeleton of Mauligobius maderensis, male, 87.0+20.5 mm, Madeira. (a). suspensorium, lateral view; (b) preopercular bridge to symplectic (right); (c), glossohyal, dorsal view; (d), urohyal, lateral view; (e) pectoral girdle, medial view; (0, scapula (right); (g), caudal skeleton, lateral view. Art, articular: d, dentary; h, hyomandibular, mpt, metapterygoid; mx, maxilla; op, operculum; pal, palatine; pmp, premaxillary pedicel; pmx, premaxilla; ptg, pterygoid (ectopterygoid); q, quadrate; sop, suboperculum; sy, symplectic; x, preopercular bridge to symplectic; cor. coracoid; cl, cleithrum; epu, epural polate; hsp, haemal spine; hyp I , parhypural I ; hyp 2 + 3. lower fan hypural; hyp 4+5 , hypural plate; hyp 6, hypural 6; nsp, neural spine; pop, preoperculum; pts and pti. upper and lower arm of post-temporal; r 1-4, pectoral radials; sc, scapula; scl, supracleithrum; v 28 h, haemal spine of last vertebra.

GOBllD FISHES OF T E M P E R A T E M A C A R O N E S l A 387

caniniform; tongue truncate or rounded; branchous tegal membrane attached along entire lateral margin of isthmus, from immediately anterior to pectoral lobe; gill-rakers simple on first arch, spinulose on rest.

Cephalic lateral-line system (Fig. 10) with anterior and posterior oculoscapular and preopercular canals, carrying pores 0, A, K , w , a, j?, p, p l , p2 and 7, 6, E , respectively (with pores p and p1 becoming contiguous; sensory papillae (G. maderensis) in seven transverse suborbital rows; row I vertical; row 2 well separated from, but row 3 near, lower border of orbit; row 5 divided by row b into upper and lower segments, latter of equal length but below upper segment of 6; lower segment of row 6 extending by a few papillae below level of row d; row b ends anteriorly behind row 4, and posteriorly distant from pore 6; anterior oculoscapular row XI ends anteriorly well behind pore j?, opposite row z ; anterior dorsal row u remote from fellow in dorsal midline; row g not anterior to lateral ends of row 0; snout with four median preorbital rows; no interorbital papillae; body with single median trunk series.

Skeleton (Fig. 1 1 ) with diagnostic features of subfamily Gobiinae given by Miller (1973b); neurocranium with postorbital region broad, longer than orbital and preorbital regions com- bined; interorbital septum narrow; basioccipital in contact with prootic; intercalar large, related to basioccipital, prootic and pterotic, leaving small opisthotic fenestra; supraoccipital separating epiotics in midline, not meeting sphenotic below frontal, and with crest attenu- ated posteriorly; mesethmoid with concave anterior edge meeting processes of vomer reflected over ethrnoidal cartilage; no vertical mesethmoid ossification in anterior corner of orbit; prefrontal present; hyomandibular with two articular struts to sphenotic and pterotic; metapterygoid extending from hyomandibular to quadrate with bridge to latter enclosing foramen; preoperculum lacking spines, with narrow process meeting rear comer of symplectic below lower edge of hyomandibular; single pterygoid (ectopterygoid) related to lower half of palatine shaft; maxilla without dorsal strut to fellow of opposite side; premaxilla with triangular crest, pedicle not reaching back to interorbit; maxilla with subacute distal end; glossohyal fan-shaped, with truncate anterior edge; urohyal with concave rear edge, lower corner attenuate; basibranchials ossified; lower pharyngeal plates separate; pectoral girdle with vestigial scapula, inverted V-shaped, and carocoid related to lowest radial; vertebrae 28 (including urostyle); interspinous bones of second dorsal and anal fins not doubled between neural and haemal spines; caudal skeleton with one epural.

Named for Mr G. E. Maul, Funchal, in recognition of his help in providing material and information used in work on Macaronesian gobies.

In addition to G. maderensis, study of the type of the West African Gobius nigri Giinther, I86 I , has suggested that this infrequently reported species may be provisionally placed in Mauligobius. The type, a male, 5 4 + 12.5 mm (BMNH 1847.4.4, sp. 45.850), from “the Niger Expedition”, is described in Appendix 11. Basic similarity with G. maderensis may be noted in head and cheek squarnation, pelvic disc shape, development of free pectoral rays, and anterior nasal tentacle, but the specimen shows more extreme development of trends by which G. maderensis differs in these features from related Gobius species. Thus, in G. nigri, all the cheek is covered with scales, half of all the pectoral rays are free from membrane, the pelvic disc is very short and rounded, with large lateral lobes, and the anterior nasal tentacle is multifid. Unfortunately, patterns of head sensory papillae are not well preserved, but a transverse arrangement of suborbital rows is evident and head canals resemble those of G. maderensis.

388 P. J . MILLER

The two species included in Mauligobius may be separated as follows. ( 1 ) Body subcylindrical; nape subhorizontal; mouth oblique; LL 50-57; D2 1/13; A 1/11-12; lower areas of both cheek and opercle always naked; anterior nostril tentacle usually simple. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . maderensis Valenciennes, 1837. - Body flattened ventrally; nape very oblique; mouth horizontal; LL 42-44; D2 I / 10; A 1/8; lower areas of both cheek and opercle scaled; anterior nostril with multifid tentacle. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . nigri Gunther, 186 I .

Mauligobius maderensis (Valenciennes, I83 7) (Fig. 12; Plate I )

FIG. 12. Muuligohius muderensis. male, 97.0+24.5 mm (MMF 2 1481), Baixa Larga, Madeira.

Gobius maderensis Valenciennes, 1837, 12: 55 (Madeira). (?) Gobius niger L., p: Lowe, 1839: 84; 1849: 9. (?) Gobius niger L., Lowe, 1841: 185. Gobius niger L: Jordan & Gunn, 1899: 346 (not Linnaeus, 1758). Gobius paganellus L. (part): Steindachner, 1868: 4 14, 4 15; Collett, 1890:

Linnaeus, 1758). Gobius maderensis: Giinther, 186 1 : 57; Von Martens, 1876: 39 1 ; Vinciguerra,

Zugmayer, 191 1: 113; Metzelaar, 1919: 281; Brito & Lozano, 1980: 175. Bathygobius paganellus: Fowler, 1936: 1000, 1003, 13 16 (part). Zosterisessor maderensis: Miller, 1973a: 5 14 (Clofnam no. 162. 26. 2). Mauligobius maderensis: Miller, 198 I : 8.

222 (not

893: 321;

Mater&zl. MADEIRA: 2 males, 122+29 and 109 + 2 8 3 mm, and another, 125+29 mm (MNHN A1 160), and 2 males, 105+25 and 90+23 mm, and one female, 107+22.5 mm (MNHN Al164), probably syntypes of Gobius maderensis; 2 males, 68.5+d and 122+31.5 mm and one female, 87.0+22.0 mm (BMNH 99.1.16.1 1-16), A. J. Arundell; 10 males, 36 + 10-75 + 19.5 mm, 6 females, 29 + 8-48 + I3 mm, and 23 of undetermined sex, 2 1 .O + 6 . 0 4 2 + 1 1-5 mm (BMNH 1953.1 1.1.300-387, part) from Baia do Salto do Cavalo, Funchal, 24 August 1953, coll. D. W. Tucker; 3 males, 32 .0+9348 .5 + 12.5 mm, 2 females,

PLATE 1. Mauhgohrut madcven,rc. from Madeira: (a) male, 44 0 + I I .0 mm: (b) female, 8 I .O + 18.5 mm; (c) male, 97.0+24.5 mm (MMF 21481): (d) female, I14,0+27,5 mm. Scalc (a) 5 mm, (bHd) 10mm.

GOBllD FISHES OF TEMPERATE MACARONESIA 39 I

40.5 +9.5 and 8 1.5 +20 mm, and 3, sex indeterminate, 28.0-t 7.5-37.0+9.0 mm (BMNH 1953. I 1. I . 388-394), from Santa Cruz, August and September 1953, coll. D. W. Tucker, F. Escorcio, E. Silva; 2 males, 97+24.5 (MMF21481) and 106+26 mm (MMF21484) and 3 females, 71 + 15.5-76+20 mm (MMF 21480, 21482, 21483) from Baixa Larga, 29 May 1965; 5 males, 66+ 14.5-120+27.5 mm, 6 females 45.0+ 12.0-1 14+27.5 mm and 2 of undetermined sex, 2 3 3 + 6.5 and 24.0+ 7.0 mm, from Barreirinha, c. 0.5 km W. Funchal, 26 & 27 July 1975, coll. A. C. Wheeler; 17 males, 24.5 + 7.0-66.0 + 17.0 mm, 17 females, 23.0+7.0-91+22 mm, and 7 of undetermined sex, 18.0+5.5-29.0+8.5 mm, from Ponta Gorda, c. 2 km W. Funchal, 25 July 1975, coll. A. C. Wheeler; one male, 69+ 18 mm, Pta doesReisMagos,Canico,c. 8.5 km E. Funchal, 30 July 1975, coll. A. C. Wheeler.

SALVAGE ISLANDS: 4 males, 76.0+ 18-5-13.0+25.0 mm and 3 females, 73.0+ 184-100.0+27.5 mm (BMNH 95.5.28.1 15-121), Great Salvage, coll. Baring & Grant; 2 males, 37.5 + d and 1 18 + 32 mm (MMF 143 1 9 , 18 July 1958; one female, 37.5 + d mm, and 2 ofundetermined sex, 33-0 + 7.0 and 36.0 + 9.0 mm.

CANARIES: one male, 68.0+ 18.5 mm (NHMW 30783), det. Steindachner; one male, 83-0+ 19.0 mm, and one female, 88.0+2 1-0 mm, from Mogan, Gran Canaria, 8 April 1977, coll. J. K. Dooley; 3 males, 38.0+ I1-79.5+ 19 mm, and 3 females, 5 8 . 5 + 16- 83.0+20.0 mm (MHNG 1571.21-73) from Los Cristianos, Tenerife, 29 December 1977, coll. P. Haymoz.

General description. Body proportions given in Table 1V; other features as genus; body subcylindrical, laterally compressed towards tail; head postorbital profile slightly convex, first dorsal origin above eye level; snout oblique, usually somewhat shorter than eye- diameter; anterior nostril tube (Fig. I3 (a)-(f)) with process from posterior rim, usually an elongated tentacle, about equal tube length, rarely bifid, or even multifid (five processes), to above labial groove but shorter in juveniles. Eyes moderately large; mouth oblique, with posterior angle below anterior half of eye; tongue truncate.

Fins. D1 VI (VI: 72 observations); D2 1/13 (13-14; 13: 69, 14: 3); A 111 1 ( 1 1-12; 11: 57, 12: 1 1 ) (terminal bifid ray of D2 and A counted as one); (branched rays) 14-15; P 19-20

Fin bases and lengths as percentage of body length given in Table IV. As percentages of standard length, in male of 87+20-5 mm, DI rays were I: 13.2, 11: 15.5, 111: 16.3, IV: 15.5, V: 14.9, VI: 12.1; D2 rays I: 13.8, 1: 15.5, 6: 17.2, penultimate: 16.7, last: 14.7; A rays I: 8.1, I : 11.0, 5: 14-4, penultimate: 16.1, last 13.8.

First dorsal fin arises behind upper origin of pectoral fin; last ray above rear third of pectoral fin, fin profile moderately low (Fig. 13(g)); dorsal spinous rays not markedly free at tips; when depressed, in males, DI IV may extend to D2 I/1, D1 V and VI to D2 1 , and, in females, D I IV to D2 I, D 1 V and VI to D2 111. Interdorsal space (Fig. 13(g)) membranous for about nine-tenths to entirety in adult males, three-fifths to almost entirety in females, total length (between DI VI and D2 I ) 5-4-8-0 (males), 7-0-7.9 (females), as a percentage of siandard length. Second dorsal fin commences above anus, with last ray opposite last anal

(18-20; 18:2,19:44,20: 19);VI/5+1/5.

FIG. 13. Muuligohius mudermsis. (a)-(fl snouts in oblique dorsal view from rear, to show variation in anterior nasal process; (a) juvenile, 2 I .O + 6.5 mm; (b) female, 63.0+ 18.0 mm; (c) female, 70.0 + 18.0 mm, (d) female, SI.O+ 18.5 mm; (e) male, 106.0+26.0 mm; (0 male, I 18.0+27.0, all from Madeira; (9) form of first dorsal and interdorsal space, in male, 120.0+27.5 mm, Madeira; (h) pectoral fin upper edge with free rays, in relation to first dorsal fin in male, 118.0+32.0 mm (MMF 143 15). Salvage Islands. Abbreviations as Fig. 3.

392 P. J . MILLER

TABLE IV

Body proportions ofMauligobius maderensis. For melhod of measuring, see Miller (1969); values given are range and, in parenlheses, mean and standard deviation

SL (rnm) n

Males 66-120

6

Females 71-1 14

9

Yo SL, H Hw SN/DI SN/D2 SN/AN SN/A SN/V CP Dlb D2b Ab CI PI VI Vd Ad Aw CPd V/AN

O/o CP, CPd

YoH, SN E PO CHd Hw

Yo E, I

Yo V/AN, V

246-27.1 14.2- 1 6.3 33.8-36. I 534-55.2 5 I .5-56.8 57.5-6 I .2 25.8-32.0 17'5-20.9 11'8-14'3 30.0-3 3.3 2 14-23.2 2 2.0-2 5.3 20.8-24'7 17.9-20. I 2 1.2-24.6 20.0-2 I . I 11.8-15.0 124-14.0 23'7-27'5

60'9-76.7

2 14-26.2 2 I .9-30.9 50'6-54.2 24.7-3 1.3 57'1-66.1

334-71.4

67.2-84.8

(25.7f 1 . 1 1) (15.5k0.96) (34.450.87) (53.6k0.79) (54.4k 1.77) (59.3k 1.49) (28.6 12.20) (18.9f 1.29) (13.1 1t0.88) (3 1.3 1t 1.37) (22.2 k0.70) (23.4k 1.26) (23.0k 1.36) (19.1 k0.81) (22.3 k 1.33) (20.6 f 0.4 1) (12.9f1.23) (1 3.3 f 0.62) (26.3 k 1.49)

(70.5 k 7.36)

(23.2 k2.10) (27.6 k 3.20) (52.1 1t 1.55) (28. I 1t 2.55) (60.3 k 3.25)

(57.5 f 14.5)

(73.4 f 7.10)

24'2-26'4 13.9-1 5.8 32'7-35'5 5 I .9-54.9 524-56.6 586-62.3 264-3 1. I 17.3-1 9.2 1 1.0-1 3.9 30.8-3 3.5 2 I a-23.9 2 143-26.3 22.3-24.9 18.0-1 9.7 19.7-22.4 19'6-21.4 11.2-14.5 12.5-1 3.8 25.1-28.3

65.5-76.0

17427.2 2 5.6-3 3.3 48.1-53.3 224-29.3 5 5.4-62.8

2 5'9-53.3

644-78. I

(25.5f0.62) (35.0f0.65) (34.5 k0.87) (53.5 f 1.04) (54.0 1t 1.40) (60.4f 1.20) (27.8 f 1.34) (18.2 1t0.79) (12.4 k0.82) (32.0k0.86) (22.4k 1.00) (23.7 k I .43) (23.0 k 0.86) (18.9f0.67) (21.21t0.87) (20.4f0.75) (12.3f 1.08) (1 3.1 f 0.42) (26.6f 1.01)

(72.0 k 4.37)

(223 f 2.85) (30.0 f 2.69) (51.4k 1.62) (25.1 k2.21) (58.7 f 2.84)

(37. I k9.07)

(70.6 f 4.02)

ray and posterior tip reaching to or over upper origin of caudal fin, beyond vertical of midline origin of latter in some males. Anal fin commences below D2 1 or 2; posterior tip reaching four-fifths of distance to, or over, lower origin of caudal fin in males, from three-quarters to lower origin of caudal in females. Pectoral fin extends back to below origin of second dorsal; uppermost five rays (Fig. 13 (h)) branched and progressively free from membrane, first for about three quarters of length; tip of sixth more or less free in large individuals; pectoral free rays reach to at least middle of gap between pectoral origin and first ray of first dorsal fin. Pelvic disc (Fig. 14 (a) and (b)) short, rounded, distant from anus in larger fish, but ellipti- cal and reaching anus in juveniles; anterior membrane high, with midline height at least three-quarters length of pelvic spinous ray and prominent lateral lobes, one eighth to one third width of membrane between lobe tips. Caudal fin rounded.

393

Scales. Body covered with scales, ctenoid in original and replacement form; predorsal area to rear border of orbits, upper opercle, upper rear cheek (Fig. 14(c)), and breast, with small cycloid scales; LL 53-55 (48-57), 48: 1, 50: 4, 51: 7, 52: 6, 53: 23, 54: 16, 55: 9, 56: 7,

2, 23: 3, 24: 2, 25: 7, 26: 9, 27: 7, 28: 7. 29: 2, 30: 1); opercular scales 7-8 (6-1 1 ) along upper edge by 4-6 (3-9) down to papilla row 0s; cheek scales 6-7 (5-10) from upper end of vertical preopercular groove forwards to suborbital papilla rows 6, 5 or even 4, and 4-6 (2-8) downwards to row b or to between row b and d; breast scales 12-17 (9-17). In an alizarin-stained individual of standard length 29 mm, opercular scales were present but preopercular scales absent; the latter are well developed at 38 mm.

GOBllD FISHES O F T E M P E R A T E M A C A R O N E S I A

57: 4); TR 17-18 (16-21; 16: 6, 17: 20, 18: 18, 19: 2, 20: 1 , 21: 1); PD 25-28 (22-30, 22:

FIG. 14. Muuligobius mudrrrnsis. (a) and (b) Pelvic disc in male of Fig. 13(h), and in juvenile. 23.5+6.5 mm, Madeira; (c) head squamation in male, 84 .0+ 19.0, Madeira; canal pores stippled; (d) squamation at origin of caudal fin in female, 68.0+ 17.0 mm, Madeira; ctenii omitted from (c) and (d). Abbreviations a s Fig. 3.

394 P. J. M I L L E R

VertL.brae. 28 (27-28; 27: 1 , 28: 61), including urostyle; an abnormality found in one specimen involved vertebrae 24 and 25, fused, each lacking a half centrum (posterior and anterior respectively).

Coloration (Fig. 12, P1. I; Appendix I , P1. 11). Body (preserved) dark brown above, with narrow pale “saddles” (below first dorsal fin, origin and middle of second dorsal, and just to rear of second dorsal base), descending more or less obliquely forwards to lateral midline; below midline several small dark blotches varyingly confluent with dark upper sides to prod- uce several irregular dark bars or mottling in juveniles, becoming more distinct narrow lateral bars (about 12) with increase in body size; abdomen pale, without small paired spots. Head dark brown, cheek and opercle with small but distinct pale spots; dark blotch above oculoscapular furrow immediately behind eye; others in middle of anterior edge of opercle (behind pore 6 ) , in anterior lower corner of opercle, and above pectoral lobe; underside of head and breast pigmented, with three more or less evident dark spots along hypohyal ridge, and darker geniohyoid area. Dorsal fins with series of pale spots along base of interradial membrane and proximal part of fin; broad distal dark band, especially on first dorsal, and a pale upper edge, narrower on second dorsal fin. Anal fin dark, especially distally, with narrow white edge. Pectoral fin lobe with upper and lower corners dark, more or less speckled pale, and no well defined mark on fin, but middle part dark beyond pale proximal area at base. Pelvic disc dark centrally, with paler anterior membrane and periphery of disc. Caudal fin base with upper and lower pale corners, of greater extent than the adjacent dark spots on base; fin dark with vertical white striae on proximal part and very narrow pale edge. In adults of both sexes, head, body and median fins may become very dark (PI. I , (d)).

Lateral-line system (Fig. 10, 16 (b)). As in generic diagnosis and Table V; also median trunk series ltm of short transverse rows, anteriorly on every second or third scale, and then, below rear end of second dorsal base, on every lateral midline scale to end of caudal peduncle; dorsal trunk series Id, opposite middle of D1 IV, in two vertical rows (nine and 12; in 80 mm female); ventral trunk series Iv not discernible; caudal series of longitudinal rows along three caudal fin rays. Supernumerary suborbital rows have been found in a number of individuals, flanking rows 2-4 or the lower section of 5.

Biology. Mauligobius maderensis has been recorded with certainty only from the continental rise islands of Macaronesia, i.e. Madeira, the Salvages, and the Canaries. The occurrence of M. rnaderensis at the Azores, suggested by Regan (1903), has not been confirmed in recent work on intertidal fish (Arruda, 1979, 1980~; CANCAP project). The species may therefore rate as one of the few endemic shore fish of the continental rise islands, among a predominance of warm-temperate Atlantic-Mediterranean (Lusitanian) forms (Ekman, 1953; Briggs 1974), and its mode oforigin is discussed below. Given the prevailing southward current, there is the possibility that M. maderensis might be found on the neighbouring coast of southern Morocco.

This goby has been found in intertidal rock pools, usually with G. paganellus (see Appendix I), but not extending down to sublittoral rocky grounds like the latter species (Brito & Lozano, 1980). At Madeira, Mr A. C. Wheeler obtained M. maderensis (listed in Material) fro’m pools situated between MTL and HWN, in size from small basins, 1 m x 1 m x 10-30 cm deep, with dense tufts of calcareous algae, to much deeper, barer pools, 20 m x 10 m x I m, or extensive shallow flooded areas, reaching 30 m x 20 m, around boulders with considerable seaweed growth. An example described by Zugmayer (19 1 l), from Great Salvage island, was also obtained intertidally.


SL 21-28 70-84 114-120 SL 21-28 70-84 114-120 n 4 5 3 n 4 5 3

PO: rl rz .s’ J.2

c 2

CI

( ‘ 2

“I

so: I 2 3 4 5 6 7 h d

PM: P

1 1’

.Y

4 3-4 2-4 3-5 3-6 4-1 2 4-8 6-8 3-4

7-12 5-8 5-10 4-12 9-1 3

11-18 2-4

10-16 22-32

42-58 29-3 7

5-6

7-14 6-12

13-26 14-24 11-26 13-32 14-27 25-38 14-22

22-35 15-26 2 1-39 2 9-4 3 27-45 34-54

8-1 5 3 1-42 5 5-8 8

119-I64 71-102 24-39

13-19 9-1 I

17-24 21-31 19-30 21-35 24-26 27-42

8-26

3 0 4 2 17-25 32-36 33-50 48-55 52-60

7-17 32-53 52-1 18

120-20 I 97-129 25-53

0s: .Y’ Y L

‘I f r J’ u.\ I U.\?

U \ ‘

b I la la

OP or

01

AD: I I

,r

0.1

0

I??

h

9-12 4-6 4-6 2-4 2-6 3-4 8

6-9 7-12 3-4 3

ab

16-2 I 7-12 7-9

4-8 5-8 3-5 4-5 9-10

13-34 20-30 12-23 6-14 5-26 9-30

2 3-3 8 15-39 38-53

7-18 5-9 16

50-67 18-27 I 2-23

10-1 7 9-19 5-10 8-1 3

12-3 1

27-46 7-43

19-25 5-1 I 6-27

18-39 24-4 7 24-37 46-55 1 4-2 3 6-1 2 19

63-78 25-3 I 19-29

12-14 11-16 9-12

11-17 29-42

Maximum standard length recorded for males is 125 mm, and, for females, 1 14 mm, total lengths being 154 and 141.5 mm, respectively. Other aspects of phenology have yet to be investigated, although it seems likely that M. maderensis will resemble the larger Gubizrs species (C. paganellus and G. cobitis), studied by Miller (196 I ) and Gibson (1970), in later age at maturity and longer life-span than is seen in smaller epibenthic species (Miller, 1979, 1984).

A,ffinities. The genus Mauligobius, as based on Gohius maderensis, belongs among certain eastern North Atlantic and European gobiines distinguished by (i) six to eight transverse suborbital rows of sensory papillae, ( i i ) at least 28 vertebrae (including urostyle), and (iii) head and predorsal squamation, extending onto the upper opercle and, in a few cases, the cheek. As well as Mauligobius, this group comprises Gobitrs L., with about 16 Atlantic- Mediterranean species and the monotypic Zosterisessur Whitley (2. ophiucrphalus (Pallas)) found in Mediterranean and Black Sea lagoons, as well as the endemic Ponto-Caspian Neogobius Iljin (including Proterorhinuh Smitt) and b4e.sogubiu.s Bleeker, with about a dozen species altogether (Ragimov, 1967; Zambriborsch, 1968; Miller, 1973a, In press). Outgroup comparisons suggest that the head and predorsal squamation found in these genera is a synapomorphy indicating common ancestry, and that other features seen within this group, such as a vertebral count of only 28, the anterior nasal process, suborbital row 7 continuity

396 P. J . M I L L E R

of anterior dorsal rows 0, free pectoral rays, and vertical bars in body coloration, are more generalized, plesiomorphic characters. It is suggested that, following Tethys subdivision, the common ancestral stock of these modern genera lived in the mid-Miocene Mediterranean, including the vast Paratethys extension of this sea. Among modern forms, it is possible that a primary phyletic dichotomy separates Gobius and Mauligobius from the Ponto-Caspian representatives and Zosterisessor. The derived character-states denoting this separation are, in the former pair of genera, appearance of a midline space between anterior dorsal row 0,

while loss of the anterior nasal process and at least great reduction of free pectoral rays has occurred in the latter grouping. This dichotomy is probably the result of later Miocene separation of the Paratethys basin, housing the common ancestor of Zosterisessor. Neogobius and Mesogohius in the initial Sarmatic phase, from the eastern basin of the Mediterranean, whence the distribution of Gobius/Mauligobius stock would have extended into the adjacent eastern Atlantic. Later, during the Messinian desiccation crisis, which followed closure of the Mediterranean (Hsu et al., 1977), this line must have survived only in Atlantic waters.

Vicariance factors to account for the divergence of Mauligobius and Gobius, and the subsequent onset of speciation in the latter, are difficult to identify. They may have been provided by the cycles of Mediterranean reopening and closure believed to have operated over the end of the Miocene and beginning ofthe Pliocene (Hsu et al., 1977). Among these gobies, the species of Gobius are linked by what is here considered to be a synapomorphic loss of suborbital row 7. The two Mauligobius species share extensive cheek squamation, which, as in Zosterisessor and the Ponto-Caspian genera, is absent, in similar form, from all Gobius species, except G. cruentatus Gmelin and a Mediterranean variant of G. paganellus (G. punctipinnis Canestrini). However, the species of Gobius with cheek squamation differ markedly in assemblages of derived characters, comprising, in G. cruentatus, the extension of suborbital rows 2 and 3 to the orbit and of row x before pore p , moderate reduction in free pectoral rays, emargination of the pelvic disc, and reduction or loss of the anterior pelvic membrane, and, in G. paganellus, the multifid anterior nasal tentacle, divided suborbital row d, and short suborbital continuation of the anterior oculoscapular canal. Accordingly, cheek squamation in the genus Gobius may be regarded as a plesiomorphic character retained from common ancestry with Mauligobius. Among other Gobius species, G. cobitis Pallas has most features in common with M. maderensis, resembling the latter in ( i ) degree of pectoral ray freeing, (ii) short, rounded pelvic disc with pronounced lateral lobes (noted by Zugmayer, 19 1 l), (iii) single long anterior nasal tentacle, (iv) continuous row d, (v) closeness of pores p and p1, and (vi) distinct lateral blotches below the midline of the body in young individuals. However, these similarities must be viewed as synplesiomorphic, involving character-states persisting elsewhere among eastern Atlantic gobiids.

As a provisional measure, Miller (1 973a) placed “Gobius ” maderensis in Zosterisessor because of the common possession of row 7 and vertical bars in coloration, features not seen in the Mediterranean/Atlantic Gobius. After more detailed comparison, it is now possible to list a number of morphological differences between Z . ophiocephalus and M. maderensis. These are, as seen in Z . ophiocephalus, (i) relatively slight freeing of pectoral rays, (i i) lack of anterior nasal tentacle, (iii) longer pelvic disc, without lateral lobes, (iv) naked cheek, (v) well separated oculoscapular pores p and p’ , (vi) union of anterior dorsal rows 0,

and (vii) reduction of suborbital row 7 to a few papillae at pore a (Borcea, 1934; Cavinato, 1952). These differences are now believed to warrant separate generic status for the two species, in current ideas on the classification of gobies based on morphological resemblance.

COBllD FISHES OF T E M P E R A T E M A C A R O N E S I A 391

In cladistic terms, it has already been noted that the first two of the characters listed can be used to suggest common ancestry of Zosterisessor with the Ponto-Caspian Neogobius and Mesogohius. Subsequent divergence of Zostcrisessor. and its Mediterranean distribution, may have been a result of the late Messinian period of transient Mediterranean lagoons, derived from Paratethys drainage, before reflooding from the Atlantic (Hsu et al.. 1977).

Origin and relationships of the Macaronesian gobiid fauna

The Macaronesian islands are products of the volcanism associated with Atlantic sea-floor spreading. Modern opinion is that the Azores and Madeira are of Miocene formation, while the Canaries are no older than the Oligocene period (Schmincke, 1976; Uchupi et al., 1976). Although the two Canary Islands (Fuerteventura and Lanzarote) flanking the Moroccan coast are probably underlain by continental crust, all the other islands are derived from oceanic shield, and it is important to appreciate that none of the Macaronesian islands are connected by continental shelf to the adjoining African coastal waters (Rothe & Schmincke, 1968; Schmincke, 1976). Even between Fuerteventura and Cape Juby, Morocco, depths exceed 1200 m (Uchupi et al., 1976).

After their volcanic origin, the coastal waters of these islands must have received a demersal marine fauna largely as a result of pelagic dispersal from adjacent mid-Tertiary continental margins. The development and present-day composition of these shallow-water communities ringing the islands thus reflects both the source and the physical environment of the prevailing ocean currents, as well as the colonizing potential of demersal species located elsewhere. These latter attributes involve juvenile viability in the plankton (a func- tion of distance to be travelled), as well as adaptiveness and competitive edge on arrival and metamorphosis to benthic life at the islands. Faunal stability and the origin of any endemic elements will have been functions of the extent and duration of the insular habitats, as well as of hydrographic fluctuation affecting isolation and local environments. The com- plexity of island colonization is noted by McDowall (I97 1 ) in the context of a full discussion by Macarthur & Wilson (1967).

Along the Iberian and North African coasts, the offshore Canary current, a southward- flowing branch of the North Atlantic drift, maintains a warm-temperate regime around the continental rise islands, and subsequently extends down as far as Cape Verde, before meeting the tropical North Equatorial current (Fairbridge & Ichiye, 1966; Allain, 1970). Conse- quently, monthly mean sea-surface temperature fluctuates from about 17 to 22°C (Madeira) and 17.5-18.5 to 21-5-22.5"C (Canaries). The Canary current also causes gyres over the north-west African shelf and promotes a southward movement of coastal bottom waters (Summerhayes et al., 1976). Nearer mid-ocean, the Azores experience sea temperatures ranging in monthly mean from 15-17 to 22-24°C. This oceanic group, although situated on the southern boundary of the North Atlantic Drift (Sverdrup et al., 1942), seems to be influenced by a weak clockwise eddy of the PortugueseKanary current (Schott, 1942; Dietrich, 1963; Meehl, 1982).

Under these circumstances, it is not surprising that the coastal fauna of the Macaronesian islands, both on the continental rise and in mid-ocean, is of chiefly Lusitanian- Mediterranean provenance (Ekman, 1953; Briggs, 1974). In the case of the gobioid fishes, earlier findings (Miller 1969, 1974) and present results fully support this view of faunal affi- nity (Table VI). Three species (Gobius niger, G. paganellus, and Thorogobius ephzppiatus),

398 P. J . MILLER

TABLE VI Sitmmarj9 of gohiid distrihiriion in irmperuic Mac.aroni~.\iu

P, p, presence at archipelago or island

Canary Islands: Fuerteventura Comera Gran Canaria Hierro Lan za ro t e La Palma Tenerife

Salvage Islands:

Madeira (and islets):

Azores: cow0 Fayal Flores Graciosa Santa Maria SBo Jorge SBo Miguel Pic0 Terceira

? P - P - P

out of the seven definitely known from the continental rise islands, are common throughout much of the temperate eastern Atlantic-Mediterranean region, a fourth, G. gasteveni, is also known from the western English Channel, as well as Madeira and the Canaries, and G. auratus is known at widely separated Mediterranean localities and is also listed for the north western, Atlantic coast of Spain (Miller, 1 9 7 3 ~ ; 1974; Miller & El-Tawil, 1974; Iglesias, 198 I) . As a more offshore species, Lesueurigobius heferojusciatus has been found off Morocco, in addition to Madeira, and L. suerii. reported from the Canaries, is common in the Mediterranean (Maul, 197 I ; Miller, 1973a). Even the endemic Muuligobius rnaderensis is closely related to the Atlantic-Mediterranean genus Gobius (see above), although its probable congener, M. nigri, is of tropical West African occurrence.

With the velocity of the Canary Current at 25-27 cm/s (Summerhayes ef al., 1976), trans- port time for gobiid postlarvae from the Cadiz Sea area to Madeira and the Canaries may, at the slowest, be about 7-10 weeks and, at average speed, less than 5 weeks, well within


the period of development before metamorphosis in at least the larger Atlantic- Mediterranean Gobizrs species (Miller, 196 I : Ballard, 1969). The transit period would be much shorter between Moroccan coastal waters and these islands. Nevertheless, a list of only seven or eight gobiid species from the continental rise islands, with only one of these extending to the Azores, suggests that the Macaronesian gobiid fauna is greatly impoverished in comparison with that of the Mediterranean and the Atlantic coasts of western Europe, where over 50 species occur (Miller, 1 9 7 3 ~ ; In press). Before seeking natural explanations for this paucity, it should be remembered that apparent lack of many species may be due to inadequate collecting. Work with an aqualung at sublittoral poison stations on otherwise inaccessible ground, trawling and dredging with covered cod-ends, and use of a fine-meshed seine net in shallow water, are all techniques which could reveal further species. For example, three additions to the list of gobies for the Canary Islands in only the last few years (Brito & Lozano, 1980; Castillo & Brito, 1982; Brito, pers. comm.) indicate that much remains to be discovered. The widespread Mediterranean species, Gohius bucchichii Steindachner, common on inshore sands in commensalism with the anemone Anemonia sitlcata and now known from Morocco (Brownell, 1978), would seem a very likely potential colonist of at least the Canaries.

However, in geomorphology, the islands of Macaronesia are characteristically mountain- ous and volcanic, with few extensive intertidal flats, and lack broad estuaries, with highly productive brackish-water shallows, so that some gobiid species may be unable to colonize these coasts for ecological reasons. High salinity in the Macaronesian area may contribute to this effect-summer values approach maxima for surface Atlantic water, being 36-36.5%0 around the Azores and 36.5-37%0 off north-west Africa (Dietrich, 1963). A biotic reason may lie behind the surprising absence of Gohius cohitis from Madeira and the Canaries, although the species occurs on the Atlantic coast of Morocco (De Buen, 1930; Brownell, 1978). In the Atlantic, G. cohitis is of intertidal occurrence, typically somewhat higher up the shore than G. pagunellus, with which the former is usually associated in such habitats (Gibson, 1972). On the islands, M. mudeerensis occurs in similar circumstances with G. paganellus, which, unlike the endemic species, extends into the sublittoral zone (Brito & Lozano, 1980). Since M. maderensis is thought to be derived from stock present around the islands, before the arrival of more temperate species (see below), G. cobitis may have subsequently failed to colonize because of competition with the established M. muderensis.

At the Azores, only one species of goby has been reliably identified (Arruda, 1979, 1 9 8 0 ~ ) . This is the widely distributed Atlantic-Mediterranean G. pagundlus, also common on the shores of Madeira and the Canaries. A record of M . rnudercwsis by Regan (1903) has not been confirmed. Despite much scope for future collecting, the finding of merely a single gobiid species may genuinely reflect the greater difficulty encountered by Old World gobies in dis- persing westwards, from the continental shelf, to the Azores, than in reaching the much closer Madeira or the Canaries. Nevertheless, among the shore fish in general, at least 80 Mediterranean species are known from the Azores, little fewer than the total (94) from Madeira (Briggs, 1974), so that discovery ofother gobiids at the Azores could be likely. Since the Azores lie at thr) lower boundary of the North Atlantic Drift, Briggs (1974) expressed surprise that their coastal fauna should be of Lusitanian-Mediterranean derivation. How- ever, a clockwise gyre at the merging of the North Atlantic Drift and the western boundary of the Canary Current, even if varying in magnitude, might ensure the preponderance of Old World species, which would need to travel a far shorter distance than American forms.

400 P. J. MILLER

Evidence for westward flow near the Azores may be seen in charts of u-component ocean currents compiled by Meehl (1982).

An opposing factor, which may reduce the number of strictly Mediterranean species reaching Macaronesia, is the eastward direction of surface water movement from the Atlantic Cadiz Sea into the Mediterranean Alboran Sea through the Straits of Gibraltar (Wust, 196 1). Postlarvae of gobiids known so far only from the Mediterranean, and thus putative endemics, could therefore be prevented from entering Atlantic waters. All the non-endemic species definitely recorded from Macaronesia are of Lusitanian, as well as Mediterranean, occurrence, and perhaps only Mediterranean species whose benthic range also extends along Iberian and Moroccan coasts are available as colonists with access to the Canary Current. This limitation is probably more apparent than real. Further Mediterranean species continue to be discovered in the Lusitanian waters of the eastern Atlantic (Maul, 1976; Almeida & Gomes, 1978) and the importance of the Straits ofGibraltar as a boundary, for at least adult dispersal of Mediterranean species to the adjacent Atlantic, seems overestimated (Ekman, 1953; Briggs, 1974).

None of the Macaronesian islands show a high degree of endemicity within the coastal marine faunas (Briggs, 1974), a feature attributed by Briggs (1966) to extinctions caused by Pleistocene cooling. Among the gobies, only Muufigobius maderensis is confined to the continental rise islands, since the supposed Madeiran endemic cited by Briggs (1974) (as Gobius ephippiatus) has now been shown to be widely distributed along temperate Atlantic and Mediterranean seaboards (Miller, 1969, and above). If the old record of M. maderensis from the Azores (Regan, 1903) remains doubtful, there is, in fact, no species ofcoastal fish endemic to temperate Macaronesia as a whole, and thus no reason for believing that the shelf faunas of the rise and oceanic islands constitute a single zoogeographical unit, as Schilder (1956) proposed. This conclusion is also supported by the meristic studies on Gobius paganellus described above (Table 111, Fig. 8). Material of this species from the Azores shows greater similarity to Mediterranean and English Channel populations than to those of the Canaries or Madeira.

An obvious hydrographic reason for this situation is that the Azores and the continental rise islands do not lie in series along the Canary Current, which may influence the former by a westward gyre before reaching Madeira and the Canaries (see above). As might be expected, meristic features of G. paganelfus from the latter islands do show similarity, including common possession of low median fin values, significantly less than those for the coast of Morocco (Brownell, 1978). The sample from Madeira has greater resemblance to the continental populations (as represented by material from the western Mediterranean). It should be noted also that the Canary and Azores populations both have some mean values significantly peculiar to these localities. These characteristics could be the result of genetic differentiation facilitated by varying intensity of colonization, or of different environmental influences on phenotypic expression. The value of meristic features, as indicators of genetic relationship between non-migratory populations, is reduced by the possibility of develop- mental plasticity (Garside, 1970). In contrast to the situation in G. paganellus, there is greater meristic similarity between Azores, Madeiran and Canaries examples of the pomacentrid Chromis chromis than between these and populations from the Mediterranean and south- western Portugal (Arruda, I980b). As less equivocal evidence, the great correspondence in the marine coastal fish fauna of the Azores with that of the rest of temperate Macaronesia, together with the lack of common endemic species, supports the idea of parallel derivation

GOBliD FISHES OF TEMPERATE MACARONESIA 40 1

from the same outside source of supply, rather than by interchange between the oceanic and continental rise islands.

The phyletic affinities of the endemic gobiid, Mauligobius maderensis, occurring around the rise islands, have been discussed above. The modern disjunct distribution of the genus Maligobius, with M. maderensis at Madeira and the Canaries, and M. nigri in tropical west Africa, may have been initiated by long-term eastern Atlantic cooling from the later Miocene into the Pliocene (Devereaux, 1967). The rise islands would have been colonized by Mauligobius stock if the latter extended along the north-west African coast well north of its present distribution, because of the warmer conditions which obtained at the Miocene- Pliocene transition. It may be assumed (Berggren & Hollister, 1974) that the set of prevailing currents was the same as at present (Fig. 15(a)). Subsequently, with cooling, the southward shift of continental Mauligobius would have isolated the islands from further arrivals of planktonic postlarvae which could be carried only to the south of retreating mainland populations (Fig. 1 5(b)). With genetic isolation, smaller island populations and increased selection pressure due to environmental change, differentiation of M. maderensis as an island species resistant to temperate conditions could have taken place, perhaps initially around Madeira and then colonizing the Canaries, by the time of the Pleistocene glaciation cycles and more rigorous cooling of the north Atlantic, starting about 3 million years ago

( 0 ) ( b ) 9 deflrction of southword current r

probobly rrposed

Dirrction of ony recolonization by M. madsrsnrls

FIG. IS. Origin and penods in history of Mauligobius maderensis. (a) Suggested Miocene/Pliocene distribution of Mauligobius stock and colonization of rise islands; (b) Pliocene/Pleistocene division of Mauligobius stock; (c) Pleistocene glaciations; suggested isotherms from Mclntyre (1974) (in Lamb, 1977), with value as departure from modem (left) and annual range (right); (d) Interglacial and Postglacial (Modem). C, Canary Islands; M, Madeira. Symbols: arrows, ocean currents (in black, transporting Mauligobius postlarvae); hatched line, northern limit of tropical marine shelf fauna, according to Briggs (1970) (a) and Briggs (1974) (b); continuous line with terminating arrows (a) and (b), extent of distribution; stippled coastline (a) and (b), continental Mauligobius as source of postlarvae.

402 P. J . MILLER

(Berggren, 1972). Survival of M. maderensis, in its present situation, may have depended on the slightly milder sea-temperatures which are believed to have prevailed around Madeira rather than at the Canaries (Fig. 15 (c)). In this area, to judge from McIntyre (1974) (in Lamb, 1977: 21, fig. 15), sea-surface temperature at the last maximum of glaciation (c. 15,000- 17,000 years ago) departed from modern values by - 2 to - 4°C (Madeira) and - 4 to - 6°C (Canaries), suggesting an annual range from > 15-> 20" for Madeira, but < 13.5-< 1 8.5" for the Canaries. If Madeira did serve as a glacial refuge for M. maderensis, then in successive interglacial periods and in present postglacial times, with amelioration in climate, there would have been little difficulty in recolonizing the Canaries by postlarvae carried down the Canaries current (Fig. 15(d)). Briggs (1966, 1974) believed that the virtual absence of endemic species ( 1 in 99) around the Azores may be the result of extinctions during cold glacial periods, three of which in the last 300,000 years are indicated by foraminifera1 assemblages in sediment cores from east of the Azores (Corliss, 1975). According to McIntyre (1974) (in Lamb, 1977), reduction in sea temperature at these islands could have been by 4 6 ° C below present mean values (giving an annual range from about < 1 1-<2OoC), a somewhat more severe regime than around Madeira, where three endemics occur in a total of 123 species ofshore fish (Briggs, 1974).

The author is most grateful to G. E. Maul and A. C . Wheeler for the supply of much material and information. His thanks are also due to L. M. Arruda, M. L. Bauchot, A. Brito, J . K. Dooley, P. Kahsbauer, M. Linley, V. Mahnert and M. J . P. van Oijen for further specimens and data, to Dorothea Ovendon for the skilful radiography of many specimens, and to J . Rayner for help in the computation of meristic results. Advice on ocean currents was kindly furnished by C . Wunsch.

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Appendix I Identijcation of small Gobius paganellus and Mauligobius maderensis

Future work on Macaronesian gobies, especially in the field, may encounter difficulty in distinguishing between small Gobius paganellus and Mauligobius maderensis, the com- monest shore gobies and likely to be found in the same habitat. These species overlap in meristic characters, and, when juvenile, both have a simple anterior nostril process, before this becomes multifid in G. paganellus, and both lack cheek scales, which subsequently develop in M. maderensis.

However, the species can be distinguished initially by coloration, and the identity confirmed by subsequent examination of the head lateral-line system. Coloration (Plate 11)

Gobius paganellus (1 ) Dark blotches below lateral midline more or less confluent longitudinally; (2) caudal fin base mostly dark edged, with conspicuous crescentic dark marks above and

(3) both dorsals with pale banding not strongly contrasted against darker markings; (4) first dorsal fin with dark spot on last interradial and posterior membrane; ( 5 ) underside of head more or less dark, without distinct spots along the hypohyal ridge;

(6) cheek and opercle with few large pale areas, nearer edge of cheek.

below midline;

and

Mauligobius maderensis (1) Dark blotches below lateral midline cleary demarcated longitudinally, but usually

(2) caudal fin base with pale areas in upper and lower corners, larger than dark spots above

(3) both dorsals with proximal series of small, distinct pale spots; (4) first dorsal fin with broad dark distal band across entire fin; ( 5 ) three distinct dark spots along the hypohyal ridge; and

uniting above with midline blotches, to form vertical bars;

and below midline spot;

GOBl lD FISHES OF TEMPERATE MACARONESIA 407

PLATE 11. (a) Gohius pagune1lu.s. juvenile, 33.0+9.0 m m (BMNH 1953.1 I . I. 300-387, part), Madeira. (b) Mauligobius muderensis, juvenile, 33,5+9.0 m m (BMNH 1953.1 I . 1. 300-387, part), Madeira. (c) M. maderensis, juvenile, 33.5 + 9.5 mm (BMNH 1953. I I . I . 300-387, part), Madeira. Diagnostic features of coloration number as in text. Scale 5 mm.

(6) cheek and opercle sometimes uniform (Plate II(b)) but often bearing numerous pale spots (Plate II(c)).

Head lateral-line system (Fig. 16)

Gobius paganellus ( I ) Pore a at end of short suborbital extension from anterior oculoscapular canal; (2) six suborbital transverse rows (1-6), parts of two (5, 6) above level of row b; (3) suborbital row d divided into two horizontal parts; and

408 P. J . MILLER

FIG. 16. Lateral-line sensory papillae and canal-pores (cross-hatched) of head in juveniles of (a) Gobius pugunel- lus. 25.0 +7.0 mm, and (b) Muuligobius rnaderensis, 23.5+6.5 mm, both from Madeira. Abbreviations and nomenclature as Fig. 2.


(4) postorbital anterior and posterior oculoscapular canals well separated with pores p and p1 distant from each other.

Mauligobius maderensis ( I ) Pore a behind eye on course of anterior oculoscapular canal; (2) seven suborbital transverse rows (f-3, parts of 5 and 6 above row h as well as short

row 7 immediately before pore a; (3) suborbital row d continuous; and (4) postorbital anterior and posterior oculoscapular canals with only short gaps between

pores p and p1.

Appendix I 1

Mudigobius nigri (Giinther, 1861) (Plate 111) Gobius nigri Gunther, 186 1 , Cat. Fish. Brit. Mus. 3: 27 (Niger Expedition). (?) Gobius macrophtalmus Osorio, 19 1 1, Mems Mus. Bocage 3: 67 (Cape Verde Islands: Ilheo Raso). (?) Gobius macrophthalmus Osorio, 19 1 1, Mems Mus. Bocage 3: 1 (figs 2,3) (error or corrig. for G. macrophtalmus Osorio). Gobius nigri: Boulenger, 1905: 188; 19 16: 24, 34, fig. 19. Bathygobius nigri: Fowler, 1936: 1000, 1001, 1316, fig. 41 1 ; Blache, 1962: 71; Daget & Iltis, 1965: 296; Blache et al., 1970: 395, fig. 1013. (?) Gobius macrophthalmus: Fowler, 1936: 13 17; Cadenat, 196 1 : 243.

Material. ‘Niger Expedition’: one male, 54+ 12.5 mm (holotype, BMNH 1847.4.4), don. L. Frazer.

Nomenclature. Gobius macrophthalmus, described by Osorio ( I 9 1 I ) from Ilho Raso, among the Cape Verde islands, is, as that author commented, a “especie visinha do Gobius nigri”. The criteria used by Osorio to distinguish the two are trivial and the description and photographs of the holotype of G. macrophthalmus agree closely with that of G. nigri, including meristic features of D1 VI, D2 I/9 or 10, A I/6 or 7, and LL 42, the divided anterior nasal process, and the free upper rays of the pectoral fins. Unfortunately, the type of G. macrophthalmus has been destroyed (Arruda, pers. comm.).

General description. Body proportions: as O/o SL, H 29.6, Hw 18.2m SN/Dl 38.9, SN/D2 60.2, SN/AN 59.3, SN/A 66.7, SN/V 24.1, CP 22.2, D l b 13.0, D2b 22.2, Ab 21.3, CP 23.2, PI 21.3, VI 22.2, Vd 22.2, Ad 22.2, CPd 13.0, V/AN 33.3; as O/o CP, CPd 58.3; as O/o H, SN 16.3, E 28.8, PO 50.0, CHd 23.8, Hw 61.3; as Yo E, 143.5 (fleshy), 13.0 (bony interorbit); as Yo V/AN, V 66.7.

Body laterally compressed towards tail, upper profile markedly convex, highest at origin of first dorsal fin, well above eye level; ventral profile subhorizontal; head postorbital profile steep; snout oblique, shorter than eye; anterior nostril tube short, wide, with fleshy process bearing four or five tentacles, longer than tube, but not reaching labial groove (Fig. 17); posterior nostril pore-like, near edge of orbit; mouth subhorizontal, with posterior angle below rear pupil and upper jaw slightly in advance; upper lip width two-thirds lateral preorbital area; tongue rounded.

Fins. D1 VI, D2 1/10, A I/8, C and P not counted, since rays difficult to separate without damage to specimen, V I/5 + I/5. Fin bases and lengths, as percentages of SL, given above. First dorsal fin arises over proximal third of pectoral fin, at apex of body profile, last ray

410 P. J . M I L L E R

Pi.ATt 111. Mudrgobrm nigrr (a) Holotype, male, 54.0+ 12.5 mm (BMNH 1847.4.4). (b) Head and pectoral fin of holotype; note cheek scales and pectoral free rays. (c) Pelvic disc of holotype; note rounded outline, much branched pelvic rays, and prominent lateral lobes of anterior membrane. Scale 5 mm.

GOBllD FISHES OF TEMPERATE MACARONESIA 41 I

above rear third of pectoral fin; when depressed, none of D1 rays reach D2 1. Interdorsal space about three-fifths membranous. Second dorsal fin commences above anus, with last ray behind vertical of last anal ray; posterior tip extending about half-way to upper origin of caudal fin. Anal fin commences below D2 2, last ray below D2 819; posterior tip reaching about half-way to lower origin of caudal fin. Pectoral fin (Plate III(b)) extends back almost to origin of second dorsal fin; uppermost ten rays short, much divided, mostly free from membrane, so that upper outline of fin concave; free rays well separated from first dorsal base; lowermost three or four rays also short with thickened edges. Pelvic disc (Plate IIl(c)) short, rounded, with short, much-branched rays; anterior membrane high, with very pronounced lateral lobes, about one third width of membrane between lobe tips. Caudal fin rounded.

Teeth. As generic diagnosis, but no lateral canine teeth in lowerjaw. Scales. Body covered with ctenoid scales, extending onto proximal fifth of caudal fin;

predorsal area, including rear two-thirds of interorbit, all opercle, and cheek forwards to vertical of jaw angle, and breast, all scaled, predorsal and opercle ctenoid, cheek cycloid. Scales in lateral series about 38-40, plus 4-6 on caudal fin, overall probably 42-44; in transverse series 17; predorsal 32; opercular 10-1 1 (along upper edge): cheek about 20 in longitudinal series; breast about 10.

A N UL

\

I S O rnrn U

( a )

0.5 m m

FIG. 17. Mair/igohiii\ m,rri. (a) snout of holotype in oblique dorsal view from rear; (b) and (c) appendages of left and right anterior nostrils respectively. Abbreviations as Fig. 3.

Vertebrae. 28 (including urostyle). Coloration (Plate 111). The holotype, after about 140 years in preservative, is now pale

brown, with an oblique dark stripe across the base of the free pectoral rays; the anterior nostril processes are noticeably dark. Giinther ( 1 86 I ) found the same “fine” specimen to be “chestnut-brown: dorsal, caudal and pectoral brown-spotted: anal and ventral blackish”.

Lateral-line system. Head canals and pores as in generic diagnosis; otherwise sensory papillae rows are abraded, except for traces ofsuborbital row 1.

412 P. J . M I L L E R

Biology. Giinther ( I 86 1) noted that the single example of Gobius nigri was obtained on the “Niger Expedition” and came into the possession of the British Museum “from Mr Frazer’s collection”. Unfortunately, the precise type-locality and habitat appear to have gone unre- corded. Boulenger (19 16), somewhat misleadingly, quotes the type-locality as merely “Niger”, suggesting a freshwater origin, from the major West African river of that name or an inland location in Niger, then a region of French West Africa to the north of Nigeria. The type specimen was collected by Mr Lewis Frazer on a British naval expedition which, in 1841, achieved only a limited penetration of the Niger delta before disease compelled withdrawal (Allen & Thomson, 1848). Mr A. C. Wheeler (pers. comm.) observed to the author that most of the British Museum fishes from this expedition are marine species and suggested that Frazer may have obtained the specimen of G. nigri at Fernando Po, used for recuperation, and the locality given for most of the birds collected on the expedition (Allen & Thomson, 1848). All the fishing mentioned by Allen & Thomson ( 1 848) in their narrative was done under marine or estuarine conditions, named localities ranging from the Cape Verde islands to Annobon (Pagalu), and including Fernando Po, as well as the Niger and Cameroon estuaries and Sao Tome. However, in an appendix dealing with animals, only two freshwater fish species are listed. As noted above, G. rnacrophthalrnus Osorio, from Ilheo Raso, Cape Verdes, is probably identical with G. nigri. Elsewhere, although G. nigri has been included in a number of recent faunal lists (see synonymy), there seem to be no definite records of other captures (Daget & Iltis, 1965) except that given by Boulenger (1905) for the mouth of the Rio Benito, Spanish Guinea. Nevertheless, a marine species found at the Cape Verdes, and perhaps Fernando Po, would be expected to occur throughout the tropical West African region.

From body shape, with strongly arched dorsal profile and flattened underside, and the short, circular pelvic disc, it may be supposed that M. nigri lives under conditions of strong current action. The limited evidence discussed above would seem to indicate that M. nigri encounters such conditions in the marine intertidal zone, or perhaps estuaries (Boulenger, 1905; Daget & Iltis, 1965) rather than a fast-flowing stream. The paper in which G. rnacrophthalmus was described (Osorio, 19 1 1 ) deals, otherwise, with many typically marine fishes.

The holotype of G. nigri is 66.5 mm in total length, and that of G. macrophthalmus was 87 mm (Osorio, 19 1 1). Other aspects of biology for this species are unknown.

Afinities. The reason for transferring Gobius nigri to Mauligobius have been discussed above, together with a scenario for allopatric speciation ofM. maderensis and M. nigri. How- ever, certain superficial features of M. nigri may suggest closer relationship with other West African forms. In extensive cheek squamation, surface meristics, and the probably transverse pattern of suborbital papillae rows, M. nigri resembles the small Gorogobius nigricinctus (Delais, 195 I ) , although the latter, on closer examination, is found to have only 27 vertebrae, no pore fl or posterior oculoscapular and preopercular canals, upper pectoral rays contained within fin membrane, a long pelvic disc without lateral lobes on the anterior membrane, and sharply demarcated vertical dark stripes (Miller, 1978). Among West African gobies, free pectoral rays and a short rounded pelvic disc, with conspicuous lateral lobes, are also seen in Rathygobius species, which, however, have only longitudinal suborbital papillae rows, naked cheek with curved groove above jaw angle, simple nasal lappet, and 27 vertebrae (Delais, 1951; Miller, unpubl.). There is thus no support for the idea that G. nigri be placed in Bathygobius, as was done by Fowler (1936) and Daget & lltis (1965).

Documents

The gobiid fishes of temperate Macaronesia (eastern Atlantic)