diff --git a/data/16/1B/87/161B87CDBA460A20FF54FF38FBD05910.xml b/data/16/1B/87/161B87CDBA460A20FF54FF38FBD05910.xml new file mode 100644 index 00000000000..4ce586fdfe7 --- /dev/null +++ b/data/16/1B/87/161B87CDBA460A20FF54FF38FBD05910.xml @@ -0,0 +1,907 @@ + + + +The insupportable validity of mosquito subspecies (Diptera: Culicidae) and their exclusion from culicid classification + + + +Author + +Harbach, Ralph E. +0000-0003-1384-6972 +r.harbach@nhm.ac.uk + + + +Author + +Wilkerson, Richard C. +0000-0001-6366-1357 +wilkersonr@si.edu + +text + + +Zootaxa + + +2023 + +2023-06-15 + + +5303 + + +1 + + +1 +184 + + + + +http://dx.doi.org/10.1094/PDIS-04-22-0755-PDN + +journal article +53758 +10.11646/zootaxa.5303.1.1 +55cb0aa4-25b5-43fc-b545-54697a22b641 +1175-5326 +8043342 +DE9C1F18-5CEE-4968-9991-075B977966FE + + + + + + + +Toxorhynchites +( +Lynchiella +) +haemorrhoidalis +(Fabricius) + + + + + + + +subspecies + +haemorrhoidalis +( +Fabricius, 1787 +) + +—original combination: + +Culex haemorrhoidalis + +. Distribution: +Argentina +, +Bolivia +, +Brazil +, +Colombia +, +Ecuador +, French Guiana, +Guyana +, +Paraguay +, +Peru +, +Suriname +, +Trinidad and Tobago +, +Venezuela +(updated from +Knight & Stone 1977 +). + + +subspecies + +separatus +( +Lynch Arribálzaga, 1891b +) + +—original combination: + +Megarhina +[ +sic +] +separata + +(subspecific status by Lane 1951). Distribution: +Argentina +, +Bolivia +, +Brazil +, +Paraguay +( + +Wilkerson +et al +. 2021 + +, +Nicaragua +deleted). + + +subspecies + +superbus + +( +Dyar & Knab, 1906a +)—original combination: + +Megarhinus superbus + +(subspecific status by Lane 1951). Distribution: +Belize +, +Colombia +, +Costa Rica +, +Cuba +, +Ecuador +, French Guiana, +Guatemala +, +Honduras +, +Mexico +, +Nicaragua +, +Panama +, +Suriname +, +Trinidad and Tobago +, +Venezuela +( + +Wilkerson +et al +. 2021 + +). + + + + +The taxonomic history of + +Tx +. +haemorrhoidalis + +involves four nominal species: + +Culex haemorrhoidalis +Fabricius, 1787 + +, + +Megarhina separata +Lynch Arribálzaga, 1891b + +, + +Megarhinus lynchi +Dyar & Knab, 1906a + +and + +Megarhinus superbus +Dyar & Knab, 1906a + +. + +Megarhina separata + +was considered a synonym of + +haemorrhoidalis + +as early as +Lutz & Neiva (1913) +. +Dyar (1928) +followed earlier workers in recognizing + +Megarhinus lynchi + +as a valid species; it was regarded as a questionable synonym of + +haemorrhoidalis + +by +Edwards (1932a) +and formally synonymized with + +separatus + +by +Lane (1939) +. Five years later, +Lane (1944) +treated + +haemorrhoidalis + +, + +separatus + +and + +superbus + +as separate species; however, he distinguished the last two based only on distribution: + +superbus + +in Central America and + +separatus + +in +Argentina +. It seems Lane (1951), without explanation, was unable to further support the specific rank of + +separatus + +and + +superbus + +and reduced them to subspecies of + +haemorrhoidalis + +. +Lane (1953) +, in accordance with his earlier interpretation, distinguished the two subspecies based only on distribution, and stated in a note that “We have placed + +T. superbus + +and + +T. separatus + +as subspecies of + +T. haemorrhoidalis + +. Such a course taken by us is strengthened by the fact that the zoogeographical distribution of the three forms is quite distinct.” This is not clear, but we believe Lane intended to say that the distributions of each of the three forms are distinct from one another. Except for +Vargas (1953) +, who either disagreed with or was unaware of Lane (1951, 1953) and recognized + +superbus + +as a species in Central America, + +separatus + +and + +superbus + +have continued to be recognized as subspecies to this day. + + +Dyar & Knab (1906a) +, in a discussion explaining why they gave the new name + +lynchi + +to the species in +Argentina +previously identified as + +haemorrhoidalis + +by +Lynch Arribálzaga (1891b) +, +Theobald (1901a) +, +Giles (1902) +and +Blanchard (1905) +, stated that “Great confusion has been caused by basing the diagnosis on the tarsal markings without reference to sex. We find that when the sexual differences are considered the tarsal markings are a useful guide in the diagnosis of the species and are a much more constant character than has been supposed.” It is noteworthy that Dyar & Knab treated + +separatus + +, also originally described from +Argentina +, as a synonym of + +haemorrhoidalis + +, and described + +superbus + +as a new species based on specimens from +Trinidad +and +Mexico +. Their concept of + +superbus + +also included the identification of + +haemorrhoidalis + +by Williston (1900) based on specimens from +Cuba +, French Guiana and +Mexico +, as well as the identification of + +Megarhinus violaceus + +by +Dyar & Knab (1906a) +and +Coquillett (1906) +based on specimens collected in Central America. +Dyar (1928) +subsequently separated + +lynchi + +and + +superbus + +from + +haemorrhoidalis + +based on the presence or absence of a basal pale band on hindtarsomere 2, present in + +haemorrhoidalis + +and absent in the other two. He distinguished + +lynchi + +and + +superbus + +based on environmental location: + +superbus + +“From the northern edge of the tropics” and + +lynchi + +“From the southern edge of the tropics”. These distinctions were restated, using slightly different terminology, by +Lane (1944 +, +1953 +), an indication that he either accepted the observations of Dyar without further study or he was unable to find additional characters to distinguish the three nominal forms. + + +In his treatment of the + +Toxorhynchites + +(as + +Megarhinus + +) of “ +Brasil +Meridional” (southern +Brazil +), +Lane (1944) +stated the following (translated from the Portuguese). + + +It is very interesting to note that, while this species [ + +haemorrhoidalis + +] occurs in the Guianas and the Amazon Valley, the two related species are found, one in the North ( + +superbus + +) and the other in the South ( + +separatus + +). The distinguishing characteristics of these three species reside in the development of the abdominal tufts of the males and in the marking of the tarsi of the females. + + +We think it very likely that they represent a single species and that both + +superbus + +and + +separatus + +are just geographic forms. A definitive solution of this case is impossible for us due to lack of material. + + +The available descriptions of the leg markings of both sexes are confusing and lack explicit detail, but it appears that they are the same in all four nominal forms except for hindtarsomere 2 of females, which is pale basally in + +haemorrhoidalis + +and completely dark-scaled in + +lynchi + +, + +separatus + +and + +superbus + +. The lateral tufts on the posterior abdominal segments of males are said to be more strongly developed in + +haemorrhoidalis + +than they are in the other three nominal forms, but the degree of development has not been made explicit. +Dyar & Knab (1906a) +indicated that strictly red tufts are only present on abdominal segment VII in + +superbus + +but are present on segments VI and VII in + +haemorrhoidalis + +and + +lynchi + +. In contrast, +Dyar (1928) +stated that the male of + +haemorrhoidalis + +has “Abdominal red tufts on the last four segments”, and this is quoted verbatim by +Lane (1953) +. In their identification keys, +Dyar (1928) +and +Lane (1953) +merely indicated that the “abdominal red tufts [are] well developed” in + +haemorrhoidalis + +and are “less developed” in + +lynchi + +( + +separatus + +of Lane) and + +superbus + +. It is interesting to note that +Dyar & Knab (1906a) +distinguished the males of + +haemorrhoidalis + +and + +lynchi + +based on the length of “segments 3 and 4” [palpomeres 3 and 4] of the maxillary palpus—equal in length in + +haemorrhoidalis + +and 3 longer than +4 in + +lynchi + +. Considering what is now known about the development of the maxillary palpus of mosquitoes ( +Harbach & Kitching 1998 +), “segments 3 and 4” are actually palpomeres 4 and 5. Oddly, this character was not mentioned in later works, and the descriptions of the maxillary palpi provided by +Dyar (1928) +and +Lane (1953) +are ambiguous. Dyar stated that + +haemorrhoidalis + +has “Palpi with the third joint long and pointed” (quoted verbatim by Lane), which surely must refer to the terminal palpomere, and Lane added that + +superbus + +has the “Last palpal segment long and acuminate” and + +separatus + +is “Similar to + +T. haemorrhoidalis superbus + +.” + + +As revealed by + +da +Costa +Lima +(1931) + +, +Dyar (1928) +failed to notice that +Goeldi (1905) +had described the egg, larva and pupa of + +separatus + +, and provided a color illustration of the adult male. It is interesting to note that hindtarsomere 2 is pale basally in the male illustrated by Goeldi, indicating that the species he described is not + +separatus + +, nor any of the other three nominal forms under discussion here, all of which have hindtarsomere 2 completely dark-scaled. +Séguy (1950) +provided a similar color illustration, which seems to correctly depict the male of + +haemorrhoidalis + +. + + +The larva and pupa of + +separatus + +were briefly described by +Forattini & Lane (1952) +based on a single larva captured in the Serra do Diabo region in the western area of +São Paulo State +in southern +Brazil +that was reared to an adult. The sex of the adult was not mentioned, but it is presumed to have been a female otherwise the specimen could not have been identified as + +separatus + +. The descriptions (in Portuguese) were repeated (in English) by +Lane (1953) +. +Dyar (1928) +and +Lane (1953) +provided brief descriptions of the larva of + +separatus + +. Their descriptions lack comparable information except the former author reported that the siphon is “over four times as long as wide” whereas the latter author described the siphon as being “three and a half times basal width.” +Lane (1953) +also provided brief descriptions of the larva and pupa of the nominotypical form. He unintelligibly characterized the larval siphon as “slightly more than one time as broad as wide.” However, judging from his illustration of the terminal abdominal segments, he obviously meant to say the siphon is slightly longer than broad. + +Vargas +(1953) + +, in a key for the identification of larvae of species of + +Toxorhynchites + +(as + +Megarhinus + +) known to occur in +Venezuela +, characterized the larval siphon of + +superbus + +as being twice as long as the saddle of segment X (transliterated from the Spanish). In comparison, +Lane (1953) +described the length of the siphon of + +superbus + +as being “two and a half times greatest width.” Although these authors expressed siphon length in different ways, the degree of actual difference seems to be greater than expected for individuals of the same species. + + +The subgenus + +Lynchiella +Lahille, 1904 + +, to which + +haemorrhoidalis + +and 16 other species belong, is predominantly Neotropical, with an extension into eastern areas of the +United States +and southeastern +Canada +represented by + +Tx. rutilus +(Coquillett, 1896) + +(see below). For the most part, the current taxonomy of + +Lynchiella + +dates back to +Lane (1953) +and +Vargas (1953) +. Their studies were based almost entirely on adult mosquitoes, and the immature stages, as noted above, were described very superficially. With the exception of + +Tx. gerbergi +Belkin, 1977 + +, + +Tx. guadeloupensis + +( +Dyar & Knab, 1906a +) and + +Tx. portoricensis + +(von R̂der, 1885), the larva and pupa of which were fully described and illustrated by +Belkin (1977) +, + +Augier +et al +. (2003) + +and + +Belkin +et al +. (1970) + +, respectively, the complete larval and pupal chaetotaxy has not been studied for any other species of the subgenus. As noted by Belkin +et al +., + +Toxorhynchites + +“is amazingly similar in all [life] stages throughout its nearly worldwide distribution and great difficulty is experienced in identifying species. New World species have been diagnosed largely on the basis of light markings of the tarsi, which frequently differ in the 2 sexes and are not always reliable. The metallic coloration of the thoracic scales shows considerable differences among species but is subject to some variation and is difficult to describe accurately owing to marked changes in color depending on the angle of observation. Few specific differences have been noted in the male genitalia. To date no reliable characters have been found to separate any of the species of a group in the larval and pupal stages….”, which is obviously due to the fact that they have not been studied in comparative detail. It is worth noting that Belkin +et al +. provisionally applied the name + +superbus + +to the species in +Cuba +. Likewise, Belkin and his colleagues applied the name + +superbus + +to specimens reared from collections made in +Colombia +( +Heinemann & Belkin 1978c +); +Costa Rica +( +Heinemann & Belkin 1977a +); +Guatemala +, +Honduras +and +Nicaragua +( +Heinemann & Belkin 1977b +); +Mexico +( +Heinemann & Belkin 1977c +); +Panama +( +Heinemann & Belkin 1978a +); +Trinidad +(the +type +locality of + +superbus + +) ( + +Heinemann +et al +. 1980 + +); and +Venezuela +( +Heinemann & Belkin 1978b +; +Navarro 1996 +). Those researchers identified + +haemorrhoidalis +sensu stricto + +in collections made in northern +Brazil +and +Ecuador +(Heinemann & Belkin 1979); +Colombia +( +Heinemann & Belkin 1978c +); and also French Guiana, +Guyana +and +Surinam +( +Heinemann & Belkin 1978b +). It is certain that + +haemorrhoidalis +sensu stricto + +and + +superbus + +are sympatric in at least +Venezuela +( + +Navarro +et al +. 2007 + +). It is interesting to note that the collections made in +Colombia +, French Guiana, +Guyana +and +Surinam +also included a species identified as sp. D, near + +superbus + +. + + +From the foregoing, it should be evident that the three subspecific forms may prove to be morphologically distinct, particularly in the larval stage; there is evidence that the distributions of + +haemorrhoidalis +sensu stricto + +and + +superbus + +overlap in northern countries of South America; and available information indicates that + +separatus + +may be geographically separated from + +haemorrhoidalis +sensu stricto + +and is restricted to areas southward of approximately latitude +20° south +. In view of these indicators, we believe it is likely that + +haemorrhoidalis + +, + +separatus + +and + +superbus + +are separate species; thus, we here formally return + +separatus + +and + +superbus + +to their original specific status: + + +Toxorhynchites +( +Lynchiella +) +separatus +( +Lynch Arribálzaga, 1891b +) + + +and + + +Toxorhynchites +( +Lynchiella +) +superbus + +( +Dyar & Knab, 1906a +) + +. We anticipate that molecular data will confirm these two forms and + +haemorrhoidalis + +are three separate species. + +Toxorhynchites separatus + +and + +Tx. superbus + +are currently listed as species in the Encyclopedia of Life. Based on available morphological data and having +type +localities in +Argentina +, + +Megarhinus lynchi +Dyar & Knab, 1906a + +is retained as a synonym of + +Tx. separatus +( +Lynch Arribálzaga, 1891b +) + +. + + + + \ No newline at end of file