Tmarus

This article is about the genus of crab spiders. For the greek mountain, see Tomaros.

Tmarus is a genus of crab spiders in the family Thomisidae. The genus was established by Eugène Simon in 1875, with Tmarus piger (Walckenaer, 1802) as the type species.^[1]

Tmarus
T. africanus from South Africa
T. piger from Belgium
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Subphylum:	Chelicerata
Class:	Arachnida
Order:	Araneae
Infraorder:	Araneomorphae
Family:	Thomisidae
Genus:	Tmarus Simon, 1875
Type species
Aranea pigra Walckenaer, 1802
Diversity
> 230 species

The genus includes more than 230 described species distributed across all continents except Antarctica.

Taxonomy

The genus Tmarus has undergone several taxonomic revisions. The generic name Martus Mello-Leitão, 1943 was considered a nomen nudum and synonymized with Tmarus. Additionally, Peritraeus Simon, 1895 was found to be a junior synonym of Tmarus.^[1]

Description

See also: Glossary of spider terms

Tmarus spiders are small to moderate-sized crab spiders with body lengths ranging from 3.7 to 7.3 millimeters. Females are typically larger than males, with males being slightly smaller in size. The spiders display cryptic coloration ranging from cream to dull grey, often mottled with cream, brown or black patterns that resemble bark, providing excellent camouflage.^[2][3]

The cephalothorax is as wide as long and rather convex, declining abruptly towards the posterior border. The clypeus is porrect (projecting forward), and the ocular area is well elevated. The lateral eyes are situated on well-defined tubercles, with both eye rows appearing almost straight or only slightly recurved. The lateral eyes are larger than the median eyes, while the anterior median eyes are the smallest.^[2]

The opisthosoma often features a posterior tubercle dorsally, though some species may have an elongated body form. The abdomen is clothed with numerous short, stiff setae, which are sometimes situated on small tubercles. The legs are spinous, with legs I and II being almost the same length and longer than legs III and IV. In Tmarus species, the posterior median eyes (PME) are situated nearer to each other than to the posterior lateral eyes (PLE), which distinguishes them from the closely related genus Monaeses.^[2]

Distribution and habitat

Tmarus species are found worldwide, with the highest diversity in tropical and subtropical regions. They typically inhabit vegetation, where they hunt by ambush rather than building webs. Many species are found on bark or leaves, where their cryptic coloration provides camouflage.

Species

As of September 2025^[update], this genus includes 230 species and two subspecies.^[1]

• 
  T. stellio from Ukraine
• 
  T. cameliformis from South Africa
• 
  T. comellinii from South Africa
• 
  T. angulatus from North America

These species have articles on Wikipedia:

• Tmarus africanus Lessert, 1919 – Tanzania, Botswana, South Africa
• Tmarus angulatus (Walckenaer, 1837) – North America
• Tmarus cameliformis Millot, 1942 – Ghana, Guinea, DR Congo, Tanzania, Malawi, Botswana, South Africa, Eswatini
• Tmarus cancellatus Thorell, 1899 – Ivory Coast, Gabon, Nigeria, Cameroon, Rep. Congo, Equatorial Guinea (Bioko), Zimbabwe, South Africa
• Tmarus comellinii Garcia-Neto, 1989 – Congo to South Africa
• Tmarus floridensis Keyserling, 1884 – United States
• Tmarus foliatus Lessert, 1928 – Senegal, Ivory Coast, DR Congo, Burundi, Tanzania, Zimbabwe, South Africa, Lesotho, Comoros
• Tmarus guineensis Millot, 1942 – Guinea, South Africa
• Tmarus hystrix (Simon, 1895) – India, Sri Lanka
• Tmarus longicaudatus Millot, 1942 – Mauritania, Niger, Namibia, South Africa, Yemen, Saudi Arabia, United Arab Emirates
• Tmarus minutus Banks, 1904 – United States
• Tmarus natalensis Lessert, 1925 – South Africa
• Tmarus piger (Walckenaer, 1802) – Europe, Turkey, Caucasus, Russia (Europe to Far East), Kazakhstan, China, Korea, Japan (type species)
• Tmarus planetarius Simon, 1903 – Guinea Bissau, Sierra Leone, Ivory Coast, Nigeria, Cameroon, DR Congo, Kenya, Malawi, South Africa

Complete list of species as of October 2025[update]

Tmarus aberrans Mello-Leitão, 1944 – Brazil Tmarus aculeatus Chickering, 1950 – Panama Tmarus africanus Lessert, 1919 – Tanzania, Botswana, South Africa Tmarus albidus (L. Koch, 1876) – Australia (Queensland) Tmarus albifrons Piza, 1944 – Brazil Tmarus albisterni Mello-Leitão, 1942 – Argentina Tmarus albolineatus Keyserling, 1880 – Brazil Tmarus alticola Mello-Leitão, 1929 – Brazil Tmarus amazonicus Mello-Leitão, 1929 – Brazil Tmarus ampullatus Soares, 1943 – Brazil Tmarus angulatus (Walckenaer, 1837) – North America Tmarus angulifer Simon, 1895 – Australia (Queensland) Tmarus aporus Soares & Camargo, 1948 – Brazil Tmarus atypicus Mello-Leitão, 1929 – Brazil Tmarus australis Mello-Leitão, 1941 – Argentina Tmarus baptistai Silva-Moreira, 2010 – Brazil Tmarus bedoti Lessert, 1928 – DR Congo Tmarus berlandi Lessert, 1928 – DR Congo Tmarus bifasciatus Mello-Leitão, 1929 – Peru, Brazil Tmarus bifidipalpus Mello-Leitão, 1943 – Brazil Tmarus biocellatus Mello-Leitão, 1929 – Brazil Tmarus bisectus Piza, 1944 – Brazil Tmarus borgmeyeri Mello-Leitão, 1929 – Brazil Tmarus bucculentus Chickering, 1950 – Panama Tmarus byssinus Tang & Li, 2009 – China Tmarus caeruleus Keyserling, 1880 – Brazil Tmarus cameliformis Millot, 1942 – Ghana, Guinea, DR Congo, Tanzania, Malawi, Botswana, South Africa, Eswatini Tmarus camellinus Mello-Leitão, 1929 – Brazil Tmarus cancellatus Thorell, 1899 – Ivory Coast, Gabon, Nigeria, Cameroon, Rep. Congo, Equatorial Guinea (Bioko), Zimbabwe, South Africa T. c. congoensis Comellini, 1955 – DR Congo Tmarus candefactus Caporiacco, 1954 – French Guiana Tmarus candidissimus Mello-Leitão, 1947 – Brazil Tmarus caporiaccoi Comellini, 1955 – DR Congo Tmarus caretta Mello-Leitão, 1929 – Brazil Tmarus caxambuensis Mello-Leitão, 1929 – Brazil Tmarus cinerasceus (L. Koch, 1876) – Australia (Queensland) Tmarus cinereus Mello-Leitão, 1929 – Brazil, Guyana Tmarus circinalis Song & Chai, 1990 – China Tmarus clarus (Mello-Leitão, 1937) – Brazil Tmarus clavimanus Mello-Leitão, 1929 – Brazil Tmarus clavipes Keyserling, 1891 – Brazil Tmarus cognatus Chickering, 1950 – Panama Tmarus comellinii Garcia-Neto, 1989 – Congo to South Africa Tmarus contortus Chickering, 1950 – Panama Tmarus corruptus O. Pickard-Cambridge, 1892 – Mexico, Panama Tmarus craneae Chickering, 1966 – Trinidad Tmarus cretatus Chickering, 1965 – Panama Tmarus curvus Chickering, 1950 – Panama Tmarus decens O. Pickard-Cambridge, 1892 – Panama Tmarus decoloratus Keyserling, 1883 – Peru Tmarus decorus Chickering, 1965 – Panama Tmarus dejectus (O. Pickard-Cambridge, 1885) – India Tmarus digitatus Mello-Leitão, 1929 – Brazil Tmarus digitiformis Yang, Zhu & Song, 2005 – China Tmarus dostinikus Barrion & Litsinger, 1995 – Philippines Tmarus ehecatltocatl Jiménez, 1992 – Mexico Tmarus elongatus Mello-Leitão, 1929 – Brazil Tmarus eques Thorell, 1890 – Indonesia (Java) Tmarus espiritosantensis B. A. M. Soares & H. E. M. Soares, 1946 – Brazil Tmarus estylifer Mello-Leitão, 1929 – Brazil Tmarus fallax Mello-Leitão, 1929 – Brazil, Guyana Tmarus fanjing Yang & Yu, 2022 – China Tmarus farri Chickering, 1966 – Jamaica Tmarus fasciolatus Simon, 1906 – India Tmarus femellus Caporiacco, 1941 – Ethiopia Tmarus floridensis Keyserling, 1884 – United States Tmarus foliatus Lessert, 1928 – Senegal, Ivory Coast, DR Congo, Burundi, Tanzania, Zimbabwe, South Africa, Lesotho, Comoros Tmarus formosus Mello-Leitão, 1917 – Brazil Tmarus gajdosi Marusik & Logunov, 2002 – Mongolia Tmarus galapagosensis Baert, 2013 – Galapagos Tmarus geayi Caporiacco, 1954 – French Guiana Tmarus gladiatus Tang & Li, 2010 – China Tmarus grandis Mello-Leitão, 1929 – Brazil Tmarus guineensis Millot, 1942 – Guinea, South Africa Tmarus hastatus Tang & Li, 2009 – China Tmarus hazevensis Levy, 1973 – Israel Tmarus hirsutus Mello-Leitão, 1929 – Brazil Tmarus histrix Caporiacco, 1954 – French Guiana Tmarus hiyarensis Ileperuma Arachchi & Benjamin, 2019 – Sri Lanka Tmarus holmbergi Schiapelli & Gerschman, 1941 – Argentina Tmarus homanni Chrysanthus, 1964 – Indonesia (New Guinea) Tmarus humphreyi Chickering, 1965 – Panama Tmarus hystrix (Simon, 1895) – India, Sri Lanka Tmarus impedus Chickering, 1965 – Panama Tmarus incertus Keyserling, 1880 – Colombia Tmarus incognitus Mello-Leitão, 1929 – Brazil Tmarus ineptus O. Pickard-Cambridge, 1892 – Panama, Colombia, Bonaire Tmarus infrasigillatus Mello-Leitão, 1947 – Brazil Tmarus innotus Chickering, 1965 – Panama Tmarus innumus Chickering, 1965 – Panama Tmarus insuetus Chickering, 1966 – Trinidad Tmarus intentus O. Pickard-Cambridge, 1892 – Guatemala, Panama Tmarus interritus Keyserling, 1880 – Panama, Brazil Tmarus jabalpurensis Gajbe & Gajbe, 1999 – India Tmarus jelskii (Taczanowski, 1872) – French Guiana Tmarus jocosus O. Pickard-Cambridge, 1898 – Costa Rica Tmarus karolae Jézéquel, 1964 – Ivory Coast Tmarus komi Ono, 1996 – Japan (Ryukyu Is.) Tmarus koreanus Paik, 1973 – China, Korea Tmarus kotigeharus Tikader, 1963 – India Tmarus lanyu Zhang, Zhu & Tso, 2006 – Taiwan Tmarus lapadui Jézéquel, 1964 – Ivory Coast Tmarus latifrons Thorell, 1895 – Myanmar, Indonesia (Krakatau) Tmarus lawrencei Comellini, 1955 – DR Congo Tmarus levii Chickering, 1965 – Panama Tmarus lichenoides Mello-Leitão, 1929 – Brazil Tmarus littoralis Keyserling, 1880 – Brazil Tmarus locketi Millot, 1942 – Guinea, DR Congo T. l. djuguensis Comellini, 1955 – DR Congo Tmarus longicaudatus Millot, 1942 – Mauritania, Niger, Namibia, South Africa, Yemen, Saudi Arabia, United Arab Emirates Tmarus longipes Caporiacco, 1947 – Tanzania Tmarus longqicus Song & Zhu, 1993 – China Tmarus longus Chickering, 1965 – Panama Tmarus loriae Thorell, 1890 – Malaysia Tmarus macilentus (L. Koch, 1876) – Australia (Queensland) Tmarus maculosus Keyserling, 1880 – Colombia Tmarus makiharai Ono, 1988 – Japan (Ryukyu Is.) Tmarus malleti Lessert, 1919 – Guinea, DR Congo, Tanzania Tmarus manojkaushalyai Ileperuma Arachchi & Benjamin, 2019 – Sri Lanka Tmarus marmoreus (L. Koch, 1876) – Australia (Queensland) Tmarus menglae Song & Zhao, 1994 – China Tmarus menotus Chickering, 1966 – Jamaica Tmarus metropolitanus Mello-Leitão, 1929 – Brazil Tmarus milloti Comellini, 1955 – Cameroon, DR Congo Tmarus minensis Mello-Leitão, 1929 – Brazil Tmarus minutus Banks, 1904 – United States Tmarus misumenoides Mello-Leitão, 1927 – Brazil Tmarus montericensis Keyserling, 1880 – Peru Tmarus morosus Chickering, 1950 – Panama, Colombia Tmarus mourei Mello-Leitão, 1947 – Brazil Tmarus mundulus O. Pickard-Cambridge, 1892 – Panama Tmarus mutabilis Soares, 1944 – Brazil Tmarus natalensis Lessert, 1925 – South Africa Tmarus neocaledonicus Kritscher, 1966 – New Caledonia Tmarus nigrescens Mello-Leitão, 1929 – Brazil Tmarus nigridorsi Mello-Leitão, 1929 – Brazil Tmarus nigristernus Caporiacco, 1947 – Uganda Tmarus nigrofasciatus Mello-Leitão, 1929 – Brazil Tmarus nigroviridis Mello-Leitão, 1929 – Brazil Tmarus ningshaanensis Wang & Xi, 1998 – China Tmarus obesus Mello-Leitão, 1929 – Brazil, French Guiana Tmarus oblectator Logunov, 1992 – Russia (Far East) Tmarus obsecus Chickering, 1965 – Panama Tmarus okinawanus Tanikawa, 2024 – Japan (Ryukyu Is.) Tmarus orientalis Schenkel, 1963 – China, Korea Tmarus pallidus Mello-Leitão, 1929 – Brazil Tmarus parallelus Mello-Leitão, 1943 – Brazil Tmarus parki Chickering, 1950 – Panama Tmarus paulensis Piza, 1935 – Brazil Tmarus pauper O. Pickard-Cambridge, 1892 – Panama Tmarus perditus Mello-Leitão, 1929 – Brazil Tmarus peregrinus Chickering, 1950 – Panama Tmarus peruvianus Berland, 1913 – Peru Tmarus piger (Walckenaer, 1802) – Europe, Turkey, Caucasus, Russia (Europe to Far East), Kazakhstan, China, Korea, Japan (type species) Tmarus piochardi (Simon, 1866) – Mediterranean, Turkey, Georgia, Iran Tmarus pizai Soares, 1941 – Brazil Tmarus planetarius Simon, 1903 – Guinea Bissau, Sierra Leone, Ivory Coast, Nigeria, Cameroon, DR Congo, Kenya, Malawi, South Africa Tmarus planifrons Mello-Leitão, 1943 – Brazil Tmarus planquettei Jézéquel, 1966 – Ivory Coast Tmarus pleuronotatus Mello-Leitão, 1941 – Brazil Tmarus plurituberculatus Mello-Leitão, 1929 – Brazil Tmarus polyandrus Mello-Leitão, 1929 – Brazil Tmarus posticatus Simon, 1929 – Brazil Tmarus primitivus Mello-Leitão, 1929 – Brazil Tmarus probus Chickering, 1950 – Panama Tmarus productus Chickering, 1950 – Panama Tmarus prognathus Simon, 1929 – Brazil Tmarus projectus (L. Koch, 1876) – Australia (Queensland) Tmarus protobius Chickering, 1965 – Panama Tmarus pugnax Mello-Leitão, 1929 – Brazil Tmarus pulchripes Thorell, 1895 – Singapore Tmarus punctatissimus (Simon, 1870) – Portugal, Spain, France, Caucasus, Russia (Europe to Far East), Iran, Turkmenistan, Korea, Japan Tmarus punctatus (Nicolet, 1849) – Chile Tmarus qinlingensis Song & Wang, 1994 – China Tmarus rainbowi Mello-Leitão, 1929 – Australia (South Australia) Tmarus rarus B. A. M. Soares & H. E. M. Soares, 1946 – Brazil Tmarus riccii Caporiacco, 1941 – Ethiopia Tmarus rimosus Paik, 1973 – Russia (Urals to Far East), China, Korea, Japan Tmarus rostratus Y. C. Zhou, Qin, G. C. Zhou, Peng & Liu, 2025 – China Tmarus rubinus Chickering, 1965 – Panama Tmarus rubromaculatus Keyserling, 1880 – United States Tmarus salai Schick, 1965 – United States Tmarus schoutedeni Comellini, 1955 – DR Congo Tmarus scopullitus Liu & Zhang, 2025 – China Tmarus semiroseus Simon, 1909 – Vietnam Tmarus separatus Banks, 1898 – Panama Tmarus serratus Yang, Zhu & Song, 2005 – China Tmarus shimojanai Ono, 1997 – Japan (Ryukyu Is.) Tmarus sigillatus Chickering, 1950 – Panama Tmarus simoni Comellini, 1955 – Sierra Leone Tmarus songi Han & Zhu, 2009 – China Tmarus soricinus Simon, 1906 – India Tmarus spicatus Tang & Li, 2009 – China Tmarus spinosus Comellini, 1955 – DR Congo Tmarus staintoni (O. Pickard-Cambridge, 1873) – Spain, France, Italy, Morocco, Algeria Tmarus stellio Simon, 1875 – Southern, Central and Eastern Europe, Turkey, Caucasus, Iran Tmarus stolzmanni Keyserling, 1880 – Peru Tmarus striolatus Mello-Leitão, 1943 – Brazil Tmarus studiosus O. Pickard-Cambridge, 1892 – Panama Tmarus subqinlingensis R. Zhang & F. Zhang, 2023 – China Tmarus taibaiensis Song & Wang, 1994 – China Tmarus taishanensis Zhu & Wen, 1981 – Russia (South Siberia), China Tmarus taiwanus Ono, 1977 – China, Taiwan Tmarus tamazolinus Jiménez, 1988 – Mexico Tmarus thorelli Comellini, 1955 – DR Congo Tmarus tinctus Keyserling, 1880 – Peru Tmarus tonkinus Simon, 1909 – Vietnam Tmarus toschii Caporiacco, 1949 – Kenya Tmarus trifidus Mello-Leitão, 1929 – Brazil Tmarus trituberculatus Mello-Leitão, 1929 – Brazil Tmarus truncatus (L. Koch, 1876) – Australia (Queensland) Tmarus tuberculitibiis Caporiacco, 1940 – Ethiopia Tmarus undatus Tang & Li, 2009 – China Tmarus ungulatus Chen, Liu & Hu, 2025 – China Tmarus unicus Gertsch, 1936 – United States Tmarus vachoni Millot, 1942 – Ivory Coast Tmarus variabilis (L. Koch, 1876) – Australia (Queensland) Tmarus variatus Keyserling, 1891 – Brazil Tmarus verrucosus Mello-Leitão, 1948 – Guyana Tmarus vertumus Chickering, 1966 – Puerto Rico Tmarus vexillifer (Butler, 1876) – Mauritius (Rodriguez) Tmarus villasboasi Mello-Leitão, 1949 – Brazil Tmarus viridis Keyserling, 1880 – Peru, Brazil Tmarus viridomaculatus Ileperuma Arachchi & Benjamin, 2019 – Sri Lanka Tmarus vitusus Chickering, 1965 – Mexico, Panama Tmarus wiedenmeyeri Schenkel, 1953 – Venezuela Tmarus yaginumai Ono, 1977 – Korea, Japan Tmarus yani Yin, Peng, Gong & Kim, 2004 – China Tmarus yerohamus Levy, 1973 – Israel Tmarus yiminhensis Zhu & Wen, 1981 – China Tmarus yueluensis Y. C. Zhou, Qin, G. C. Zhou, Peng & Liu, 2025 – China Tmarus zhui Sherwood & Li, 2021 – China