SHORT COMMUNICATION

First fossil horsefly (Diptera: Tabanidae) in Miocene Mexicanamber

Jonina Strelow • Monica M. Solorzano Kraemer •

Sergio Ibanez-Bernal • Jes Rust

Received: 11 September 2012 / Accepted: 4 February 2013

� Springer-Verlag Berlin Heidelberg 2013

Abstract The fossil record of the Tabanidae is sparse when

compared with other families of Diptera. Even in amber they

are rare, probably because of their size and specific flight

behavior. Horseflies from amber are only known from Cre-

taceous age New Jersey amber as well as from the Tertiary

age Baltic and Dominican amber, but are herein described for

the first time, with Stenotabanus oleariorum sp. n., from

Mexican amber. The new species is compared to the fossil

horseflies of the same genus S. brodzinskyi Lane, Poinar and

Fairchild 1988 and S. woodruffi Lane and Fairchild 1989

from Dominican amber.

Keywords Diptera � Tabaninae �Diachlorini � Taxonomy �Taphonomy �Mexican amber

Kurzfassung Der Fossilbericht der Tabaniden ist, im

Vergleich zu anderen Dipterenfamilien insgesamt noch

sehr sparlich. Auch in Bernsteinvorkommen sind

Tabaniden selten, vermutlich wegen ihrer Große und

ihrem speziellen Flugverhalten. Tabaniden in Bernstein

sind bis jetzt nur aus dem kreidezeitlichen Bernstein von

New Jersey sowie dem tertiaren Baltischen und

Dominikanischen Bernstein bekannt. In der vorliegenden

Untersuchung wird die erste Tabanide aus dem Mexikanischen

Bernstein, Stenotabanus oleariorum sp. n, beschrieben. Die

neue Art wird mit den beiden bis heute einzigen bekannten

fossilen Tabaniden der gleichen Gattungen (S. brodzinskyi

Lane, Poinar and Fairchild 1988 und S. woodruffi

Lane and Fairchild 1989) des Dominikanischen Bernsteins

verglichen.

Schlusselworter Diptera � Tabanidae � Diachlorini �Taxonomie � Taphonomie � Mexikanischer Bernstein

Introduction

The Tabanidae or horseflies are a very large and widely

distributed family of brachycerous Diptera. Males and

females of extant species feed on nectar and other plant

sugars, while the females of some species are blood feeders

from a variety of vertebrate hosts in order to improve

ovogenesis and vitelogenesis. Some species occur across a

range of open and forested habitats (Burger 2009), while

others are confined to coastal zones. Horseflies usually rest

on foliage or on tree trunks. The females oviposit on

vegetation, commonly near aquatic or semi-aquatic habi-

tats, and most larvae are found in a variety of aquatic and

semi-aquatic habitats, usually in sand or soil of varying

wetness (e.g., freshwater, ponds and streams) where they

apparently prey on small invertebrates (Burger 2009).

Adult tabanids are, therefore, restricted to habitats with a

wet breeding site nearby. Additionally, most species of

J. Strelow (&) � M. M. Solorzano Kraemer � J. Rust

Steinmann-Institut fur Geologie, Mineralogie und Palaontologie,

Rheinische Friedrich-Wilhelms-Universitat Bonn, Nussallee 8,

53115 Bonn, Germany

e-mail: jstrelow@uni-bonn.de

M. M. Solorzano Kraemer

e-mail: Kraemer@senckenberg.de

J. Rust

e-mail: jrust@uni-bonn.de

M. M. Solorzano Kraemer

Senckenberg Forschungsinstitut und Naturmuseum,

Senckenberganlage 25, 60325 Frankfurt am Main, Germany

S. Ibanez-Bernal

Instituto de Ecologıa, A.C, Antigua Carretera a Coatepec

No. 351, El Haya, Xalapa 91070, Veracruz, Mexico

e-mail: sergio.ibanez@inecol.edu.mx

123

Palaontol Z

DOI 10.1007/s12542-013-0171-7

Tabanidae are crepuscular and only active during the

warmer period of the year on sunny days. The adult flight

activity decreases when it is cool and breezy. Indeed, in

central Amazonia the occurrence of species is almost

always restricted to the dry season (i.e., low humidity and

high temperatures) when water levels in rivers and lakes

are low (Ferreira et al. 2002). As a result of the dependency

of the Tabanidae on specific ecological conditions, it is

possible to compare living species with the fossil from

Mexican amber and to draw conclusions as to the former

amber forest ecosystem.

The fossil record of the Tabanidae contains only 16

genera and 28 species. The oldest species of a true tabanid

derive from the Cretaceous with examples reported from

China, England, Russia, Brazil and the USA (New Jersey)

(Martins-Neto and Santos 1994; Coram et al. 1995; Ren

1998; Mostovski et al. 2003; Grimaldi et al. 2011). The

family is also known from the Oligocene of France,

Switzerland and Brazil, from the Oligocene and Miocene

of the USA, from Eocene age Baltic amber, from the

Eocene/Oligocene of England, from the Oligocene of

France, Switzerland, Brazil and Colorado, from the Oli-

gocene and Pliocene of Germany, the Miocene Dominican

and Mexican age amber and from the Pliocene of Europe

and North Africa as well as from the Pleistocene of Africa

(Evenhuis 1994; Martins-Neto 2003). Tabanids are only

rarely entrapped by resin, and until now only 12 speci-

mens (8 genera and 11 species) in total have been

described from Baltic amber, Dominican amber and

African copal. The complete fossil record of all described

tabanids is listed in Table 1 (undescribed or unplaced

specimens are excluded).

In the present study, a specimen of Tabanidae

from Miocene age Mexican amber is described. The

specimen was compared to the two previously described

fossil tabanids from the Dominican amber (Stenotabanus

brodzinskyi Lane, Poinar and Fairchild 1988 and Steno-

tabanus woodruffi Lane and Fairchild 1994) as well as

with extant species of the genus from the Neotropical

region.

The comparison of the Neogene age tabanid faunae of

the Caribbean Islands with those of Central America is of

particular interest in terms of understanding the biogeo-

graphic relationships between Mexico and Hispaniola. The

Mexican amber is mined in the vicinity of Simojovel de

Allende in the state of Chiapas, Mexico. It is dated to be of

Middle Miocene age (about 20 Ma) and can, therefore, be

correlated with the Dominican amber deposits (Solorzano

Kraemer 2007). Southern Mexico is considered to be a

megadiversity region in the present day (Myers et al.

2000), and the analysis of recent and fossil insect taxa is

significant in terms of ecological, paleobiogeographical

and taphonomic studies.

Materials and methods

The horsefly is embedded in a piece of amber approximately

11 mm 9 9.5 mm 9 8 mm in size. The differentiation of

species of tabanids is based on male genital differences,

pronounced morphological differences of the head and its

appendages, thoracic and abdominal patterns and coloration,

and distinctive wing patterns. Because body colors and

pubescent coloration of the living horseflies are important

distinguishing features, it should be mentioned that the colors

of the fossil tabanids may not reflect those of the living

tabanids, but the color patterning is still preserved (as is the

case in specimens preserved in other amber inclusions). The

morphological terminology used below follows McAlpine

(1981) and Burger (2009). For the taxonomic identification

and investigation, a Leica Mz 95 and a Leica MZ 125 were

used. Drawings were rendered with the aid of a drawing tube,

and measurements are given in millimeters. Photographs of

the amber inclusion were made with a Leica MZ 16 Stereo-

microscope with a JVC ky-F70B Digital Camera. Compound

photographs merging different focus levels to a single image

were performed with using Discus software equipped with a

stacking function.

Systematic palaeontology

Family Tabanidae Latreille 1802.

Tabanii Latreille 1802, 438. Additional references:

Latreille 1802, 398; Sabrosky 1999, 296.

Subfamily Tabaninae Loew 1860.

Tabaninae Loew 1860, 14. Additional references: Philip

1947, 283 (Tabaninae Rondani 1841, 283, was used for

family level as commented by Sabrosky 1999).

Tribe Diachlorini Lutz 1909.

Diachlorinae Lutz 1909, 29 (= Diachlorini, according

with Sabrosky 1999). Additional references: Enderlein

1922, 349 (as Diachlorini); Krober 1932, 197; Philip

1941, 5, 10; Fairchild 1942, 297; Philip 1947, 284;

Mackerras 1954, 431, 439 (diagnosis); Stone 1965

(reprint 1983, 328) (Nearctic catalog); Fairchild 1969,

207 (Classification); Fairchild 1971, 36 (Neotropical

catalog); Fairchild and Burger 1994, 62 (Neotropical

catalog); Sabrosky 1999, 111 (Family-group name).

Genus Stenotabanus Lutz 1913.

Stenotabanus Lutz 1913, 487; Lutz 1914, 167 (rep-

rinted). Type-species: Tabanus taeniotes Wiedemann

(Bequaert 1924, 30). Additional references: Enderlein

1925, 354; Krober 1929, 113 (in part); Stone 1938, 31;

Philip 1941, 11; Fairchild 1942, 297; Philip 1947, 285

(Nearctic catalog); Fairchild 1969, 214 (classification);

J. Strelow et al.

123

Fairchild 1971, 44 (Neotropical catalog); Fairchild 1986

46 (Panama species); Fairchild and Burger 1994, 72

(Neotropical catalog); Chainey et al. 1999, 75 (South

American species); Burger 2009, 502, 506 (key, Central

America).

Leptotabanus Lutz and Neiva 1914, 72 (nomen nudum).

Styposelaga Enderlein 1922, 348. Type-species: Stypo-

selaga sexannulata Enderlein (orig. des. = Tabanus

incipiens Walker 1860).

Fossil species of Stenotabanus.

Stenotabanus brodzinskyi Lane, Poinar and Fairchild

1988, 594, Holotype female. Type-locality: Dominican

Republic, Santo Domingo, unspecified amber mine in

the Cordillera Septentrional mountain range, 15–20 Ma

(late Early-early Middle Miocene).

Stenotabanus woodruffi Fairchild and Lane 1989, 630,

Holotype female. Type-locality: Dominican Republic,

15–20 Ma (late Early-early Middle Miocene).

Table 1 Summarized fossil record of described Tabanidae based on Evenhuis (1994) and Martins-Neto (2003)

Europe America Asia Africa

Cretaceous Eotabanoid lordi Mostovski,

Jarzembowski and Coram 2003

(England)

Cratotabanus stenomyomorphusMartins-Neto and Santos 1994

(Brazil, South America);

Cratotabanus sp. n. Grimaldi

2011 (Brazil, South America);

Cratotabanus newjerseyensisGrimaldi 2011 (New Jersey

amber, USA)

Palaepangonius eupterus Ren 1998

(China); Eopangonius pletus Ren

1998 (China); Baissomyia reditaMostovski, Jarzembowski, and

Coram 2003 (Transbaikalia,

Russia)

Eocene Mesomyia hoffeinsorum Trojan

2002; Mesomyia stigmaticaTrojan 2002; Mesomyia cupreaTrojan 2002; Mesomyiayantarophila Trojan 2002;

Sznablomyia parvula Trojan

2002; Tabanosoma tabaniformeTrojan 2002; Pseudotabanusdereckii Trojan 2002 (all Baltic

amber)

Oligocene Tabanus vectensis Cockerell 1921

(England); Tabanus statziMoucha 1972 (Germany);

Aemodipsus bornensis Maneval

1936 (France); Chyrosops seguyiPiton 1940 (France); Hexatomaoeningensis (Heer 1864) Evenhuis

1994 (Switzerland)

Silvius merychippi Melander 1947

(USA); Tabanus parahippiCockerell 1909 (Colorado, USA);

Tabanus hipparionis Cockerell

1909 (Colorado, USA); Tabanusmerychippi Cockerell 1916

(Colorado, USA); Tabanustremembeensis Martins-Neto

Martins-Neto 2003 (Brazil, South

America)

Miocene Stenotabanus brodinzkyi Lane,

Poinar, and Fairchild 1988

(Dominican amber); Stenotabanuswoodruffi Fairchild and Lane

1989 (Dominican amber);

Stenotabanus oleariorum Strelow,

Solorzano Kraemer, Ibanez-

Bernal, and Rust 2012 (Mexican

amber)

Pliocene Tabanus sudeticus Zeller 1842

(Germany and Poland); Tabanusfossilis Grabenhorst 1985

(Germany)

TabanussudeticusZeller 1842

(Morocco)

Pleistocene HaematopotapinicolaStuckenberg

1975 (Copal

inclusion)

Undescribed or unplaced specimens are excluded

First fossil horsefly

123

Stenotabanus oleariorum Strelow, Solorzano Kraemer,

Ibanez-Bernal and Rust new species.

Type species Holotype female from the collection of the

Staatliches Museum fur Naturkunde, Schloss Rosenstein,

Stuttgart, Germany (SMNS) with inventory number Mx

245. The specimen is embedded in early Middle Miocene

amber from Simojovel de Allende, Chiapas, Mexico.

Diagnosis A small species with clear wings, appendix at

base of vein R4, with frons slightly convergent-sided

below, antennae nearly uniform colored, with basal flag-

ellomere obtuse angled, mesonotum unstriped and abdo-

men unicolor.

Etymology The specific epithet is dedicated to Hans J.

Olearius and Dr. Christian Olearius for their interest and

support of science.

Description

Female: Length of body 9.1 mm, length of wing 7.5 mm.

Fig. 1.

Head: Eyes bare, golden brown without colored trans-

verse bands. Postorbital setae long, shorter medially. Frons

narrowed below, with long dense setae at vertex and

compact basal callus (Fig. 2b). Median callus, slender,

elongated, ridge-like attaining 0.7X height of frons and

attached to basal callus. Subcallus yellowish brown, flat

and bare, ocelli inconspicuous, frontoclypeus with scat-

tered dark hairs. Antenna orange brown, porrect and scape

and pedicel with many short, dark setae, scape barley

longer than wide. Flagellum uniformly colored, basal

portion of flagellum obtuse angled, width up to 0.4 its

length, with sparse, short setae plus four more terminal

annulations covered with short strong hairs. First flagello-

mere of antennal flagellum larger than the 4 apical flag-

ellomeres. Left antenna 1.6 mm long with flagellum 2.39

longer than the length of scape and pedicel combined; basal

plate 2.49 its width, and 1.469 as long as the four annu-

lations (Fig. 2a). Palpus brown, slender, elongated with

strong setae covering the dorsal area and short and scat-

tered setae covering the ventral area. Proboscis pale brown,

length sub-equal to that of palpi but longer than height of

head; labella without sclerotizations.

Thorax: Mesonotum orange-brown without longitudinal

stripes, moderately beset with short, dark, scattered hairs

medially and longer dark hairs anteriorly. Scutellum and

notopleuron thinly clothed with long, scattered hairs.

Katatergite heavily clothed with long dark hairs. Wings

hyaline, pterostigma brown, basicosta sharply pointed

without macrosetae. Costa (C), subcosta (Sc) and R1 cov-

ered with tiny short setae (Fig. 2c). Appendix present at

base of R4, wings without distinct clouds or streaks. Legs

slender, the tibia not flattened or inflated. Tibiae dark

brown covered with short, dark hairs. Hind and fore tibiae

without spurs, mid tibiae with two strong spurs. Tarsal

claws sub-equal in size, paired on the legs where the tarsi

are preserved, tarsi missing on both mid tibiae.

Abdomen: 5.1 mm long and 1.3 mm broad with seven

segments visible, yellowish dark brown, covered with dark

setae, no color pattern preserved. Seventh tergum dark

brown, clothed with dark hairs. Terminalia of female

inconspicuous.

Remarks

The fossil horse fly described here from Mexican amber

can be assigned to the subfamily Tabaninae by the absence

of functional ocelli and also of hind tibial spurs, and to the

tribe Diachlorini by the nude basicosta, wings without

infuscate patterns, antennal flagellum with 4 annuli, and

frons slightly widened below. The inclusion in amber does

not facilitate seeing the divided condition of the ninth

tergum or the color pattern of the eyes. We have assigned

this species to the genus Stenotabanus Lutz on the basis of

the bare eyes, the width of the frons, the form of the frontal

callus, the clear wing membrane, unicolor pleura,

unstripped mesonotum, and the antennal flagellum with

obtuse angle and four annuli (Fairchild 1969).

Stenotabanus oleariorum sp. n differs from Stenotab-

anus brodzinskyi, by the presence of long erect dark setae

anterolaterally and laterally on the mesonotum, and the

lack in the latter of the appendix of R4. S. woodruffi also

lacks the appendix at the fork of R4 and differs from the

other two species by having the apical third of the wing

infuscated, the frons considerably narrower and a sharp

dorsal angle of the basal plate (Fig. 2a). S. oleariorum sp.

n. is more similar to S. brodzinskyi by the form of the

flagellar basal plate, differing by the wider frons and

antenna coloration, whereas it is more similar to S. wood-

ruffi because the frons is slender and has dull-edged angles

of the basal plate. S. woodruffi and S. oleariorum sp. n.

differ from S. brodzinskyi by being much paler in color of

the integument with only parts of mesonotum, tarsi, and

annulated part of antennae flagellum being black.

Currently, 99 species (divided into 7 subgenera) of the

genus Stenotabanus are recognized (Burger 2009). The

subgenera include: Stenotabanus Lutz 1913, Aegialomyia

Philip 1941, Brachytabanus Fairchild 1942, Cretotabanus

Faichild 1969, Melanotabanus Lutz and Neiva 1914,

Phorcotabanus Fairchild 1961, and Stenochlorops Fair-

child 1969. Of these, only Aegialomyia Philip 1941,

Brachytabanus Fairchild 1942 and Stenotabanus Lutz 1913

occur in Central America (Fairchild 1969).

J. Strelow et al.

123

Fairchild (1980) erected two groups of Stenotabanus

(Stenotabanus) based on his study on the Caribbean Island

tabanid fauna, the brunettii species group with S. parvulus

Williston 1887, S. alticolus Fairchild 1980, S. batesi

Bequaert 1940, and the fenestra group, which contains

S. fenestra Williston 1987, S. marcanoi Fairchild 1980, and

S. hispaniolae Bequaert 1940. Fairchild (1988) placed the

fossil horsefly S. brodzinskyi of the Dominican amber in

the fenestra group based on the following characteristics:

frons noticeably narrowed below and more than 4.09 as

high as basal width, hyaline wings, and the lack of con-

spicuous abdominal color patterns. In the key provided by

Fairchild (1980), S. oleariorum sp. n. resembles Steno-

tabanus marconoi Fairchild 1980 based on the narrow

Fig. 1 Stenotabanus oleariorum: 1, lateral view of entire fly; 2, lateral view of left antenna; 3, lateral view of the head and proboscis. Scale of

images 2 and 3 is 0.5 mm

First fossil horsefly

123

frons, the entirely clear wing, and the appendix at vein R4.

However, it is smaller in size (9.1 mm compared to 11 mm

in S. marconoi) and also has palpi and proboscis sub-equal

in length (whereas S. marconoi having a proboscis about

twice the length of the palpi). S. oleariorum sp. n. can be

easily distinguished from S. fenestra Williston 1987 and

S. hispaniolae Bequaert 1940 because of the blackish

appearance and black wings that the latter two species

share. In addition, S. fenestra Williston 1987 has the tibiae

and basitarsi white pillose and the basicosta and antennae

black, while S. hispaniolae has a parallel sided frons and

narrow pale sutural bands on the abdomen (Fairchild

1980).

Discussion

To date only one specimen of the genus Stenotabanus has

been described from Mexican amber, the herein described

S. oleariorum n. sp. This is the sole representative of the

family from these deposits, clearly illustrating both the

scarcity of tabanids as inclusions but also the variable

preservation potential of different insect groups. In general,

small-sized insects are more likely to become trapped in

resin, while larger and stronger insects have better chances

of escaping from fresh resin flows. Therefore, size is

probably not the important taphonomic selection factor in

the Tabanidae, because most of them are of considerable

Fig. 2 Comparison between the

two fossil tabanids from

Dominican amber with the

fossil from Mexican amber:

a antennae; b frons; c wings

(drawings from S. brodzinskyiand S. woodruffi after Fairchild

and Lane 1989)

J. Strelow et al.

123

medium to large size, (e.g., S. oleariorum n. sp.). Seasonal

factors and the extent of resin production, flight activity, as

well as other behavioral characteristics or specific prefer-

ences are probably more important factors in explaining the

poor record of horseflies in amber. This is exemplified by a

study of Bickel and Asker (2004) who collected tree trunk

invertebrate fauna in an Australian forest using sticky traps.

Not a single Tabanidae was trapped from a total of 103,504

collected insects. Even though tabanids are not tree

inhabitants they are active flyers and have, at least some-

times, the potential to get trapped in various types of tri-

angle-shaped tent and cloth traps (emergence, malaise, and

canopy traps) as well as in attractant traps (carbon dioxide

and octenol).

A comprehensive revision of the fossil record of the

Tabanidae has already been suggested by Martins-Neto

(2003), but is outside the focus of the present study

(Table 1). However, S. oleariorum n. sp. represents the

third fossil member of the tribe Diachlorini and of the

genus Stenotabanus reported in the literature. The simi-

larities in the tabanid faunas of Mexican and Dominican

amber deposits suggest a possible faunal interchange

between both regions in pre-Miocene times. Fairchild

(1969) notes that of the three tribes recognized in Taban-

inae, Diachlorini contains the most primitive members.

Furthermore, he divides the tribe into two groups: a

primitive group including Stenotabanus Lutz that is prob-

ably derived from Dasybasis and in turn may have given

rise to the more specialized groups such as Diachlorus.

This is important, since in Mexico there are more endemic

species known from the genus Stenotabanus than species

from the genus Diachlorus (Ibanez-Bernal and Coscaron

2000), but the phylogenetic relationships of the tribe and

the Neotropical tabanids in general require clarification.

The extant distribution of Stenotabanus is restricted to

the Neotropical region with reports from the southern parts

of North America, the Caribbean Islands, Central America,

and South America. The presently known Stenotabanus

fauna of Mexico consist of ten endemic species and ten

widespread species. The species of S. (Stenotabanus) that

are geographically restricted include S. abacus Philip 1954,

S. apaches Philip 1977, S. cribellum Osten Sacken 1886,

S. litotes Fairchild 1953, S. mexicanus Philip 1977,

S. pumiloides Williston 1901, S. stonei Philip 1958, and

S. subtilis Bellardi 1862, and for S. (Aegialomyia) are

S. chiapasensis Fairchild 1953, S. indotatus Ibanez-Bernal

1991, S. occidentalis Philip 1976, and S. yaquii Philip

1976. The widespread species of S. (Stenotabanus) include

S. flavidus Hine 1904, S. fulvistriatus Hine 1912, and

S. minusculus Krober 1930, and for S. (Aegialomyia) are

S. guttatulus Townsend 1893, S. jamaicenis Newstead

1909, S. littoreus Hine 1907, S. magnicallus Stone 1935,

and S. pechumani Philip 1966.These species generally

inhabit lentic habitats. On that account, the fossil Steno-

tabanus therefore indicates the existence of (temporary)

water ponds in the plain regions within the former amber

forest. As already mentioned in the introduction, female

tabanids lay their eggs on vegetation near or above aquatic

or semi-aquatic habitats, and most larvae inhabit marshes

or streams, but some larvae inhabit even dry soil such as

the immature stages of S. (Stenotabanus) incipiens Walker,

which have been found in soil near an old log (Burger

2009). If the oviposition behavior did not change, adult

species inhabited regions comparable to these environ-

ments probably within the woodlands, which would make

entrapment in resin possible.

Acknowledgments This research was possible with a postdoctoral

fellowship to M.M.S.K, no. SO894/3-1, from the German Research

Foundation (DFG). The authors would like to thank Dr. Gunter

Bechly from the Staatliches Museum fur Naturkunde Stuttgart for the

loan of the specimen. Special thanks are due to PD Dr. Torsten

Wappler (Bonn) for valuable comments and helpful discussion and

Prof. Dr. McCann (Bonn) for reading and correcting the manuscript.

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