Ingolfiellidea (Crustacea, Malacostraca, Amphipoda): a Phylogenetic and Biogeographic Analysis
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Contributions to Zoology, 72 (I) 39-72 (2003) SPB Academic Publishing bv, The Hague Ingolfiellidea (Crustacea, Malacostraca, Amphipoda): a phylogenetic and biogeographic analysis Ronald Vonk & Frederick+R. Schram Institute for Biodiversity and Ecosystem Dynamics, University ofAmsterdam, Mauritskade 57, 1092 AD Amsterdam, The Netherlands Keywords:: Ingolfiellidea, phylogeny, biogeography, new genera, paleogeographic maps, bibliography Abstract Paraleleupia Ingolfiella Additional descriptions 63 The suborder consists of 39 named Ingolfiellidea currently spe- Biogeographic analysis 65 cies. An historical overview is presented and phylogenetic and Discussion 67 biogeographic analyses are made. The result ofthe phylogenetic Acknowledgements 71 analysis the definition of new within an suggests two genera References 71 African and freshwater group, namely Paraleleupia n. gen. Proleleupia n. gen. Re-examinationof a supposedly Italian re- lict species, Metaingolfiella mirabitis, with the aid of SEM Introduction techniques reveals a half-fusion ofthe head region with the first pereionite.The issue of the function of the ‘eyelobe’ is addressed and an explanation presented after examining with SEM such The ingolftellidean amphipods are not abundant in lobes in different additional species. Furthermore, descriptions regards to the numberof species. To date, some 39 are based given onthe type-material ofMetaingolfiella mirabilis, species are recognized, a remarkably low number Trogloleleupia eggerti, Trogloleleupia leleupi, Ingolfiella lit- the wide-ranging ecological conditions toralis, considering I. tabularis, I. margaritae, I. quadridentata, and I. abyssi. An in which they occur. It is also a low number when attemptto relate the geographic distribution and ecological to other of with characteristics to the phylogeny of the Ingolfiellidea results in compared family groups amphipods a habitat paleogeographic scenario that points to a freshwater sub- partly overlapping requirements such as terranean for origin the group that dates from at least Triassic bogidiellids, with 110 species (Koenemann & Hol- times. and singer, 1999) crangonyctids (250 species, pers. comm. Koenemann). The first specimens ofIngolfiellidea were reported Contents Hansen 1903. He classified them by in in a new suborder, a rank they have continued to hold in Introduction 39 most publications ( Bousfield & Shih, 1994; Ruffo Methods and material 42 & 1989; Martin & Davis, Character Vigna Taglianti, 2001) descriptions 42 several the Cladistic despite objections over years (Dahl, 1977; analysis 57 Results Bowman & Barnard & ' 58 Abele, 1982; Karaman, Systematics It is to this small 60 1983). a challenge compare group Taxonomic diagnoses derived from cladistic analysis 60 of other well-defined crustaceans to larger groups Ingolfiellidea 60 find and try to comparable habitat preferences and Metaingolfiellidae 61 overlapping biogeographic distribution patterns. Ingolfiellidae 61 From consideration of the literature a picture Stygobarnardia 61 Trogloleleupia 61 emerges of the uniqueness of ingolfiellideans within the entire Proleleupia 61 crustacean world with regard to their Downloaded from Brill.com10/10/2021 03:55:37AM via free access 40 R. Vonk & F.R. Schram -A phylogenetic and biogeographic analysis I. World of distribution of Black dots = marine localities. = brackish conditions. Fig. map Ingolfiellidea. Open stars water Open circles = fresh water localities. incredibly diverse habitat tolerances. No other small Congo, later ranked in a separate genus: Troglole- taxon of crustaceans is found in the soft mud of leupia leleupi (Ruffo, 1951). Compared to the 2.5 well the deep-sea floor, as as in high mountain fresh- mm specimens for previous taxa this African cave in stood 14.5 maximum water river beds, or subterranean fresh, brackish lake species out at mm length. and marine interstitial waters of continental ground Then more species were discovered in Macedonia: waters and continental shelves (Fig. 1, map) Ingolfiella petkovskii S. Karaman, 1957, and I. mace- The study ofthe first species of Ingolfiella marked donica S. Karaman, 1959. S. Karaman character- this habitat diversity. Hansen (1903) described two ized their rarity when he stated that he had searched species, Ingolfiella abyssi, from 3521 m. deep sea through several thousand samples from ground water bottom out of a trawl of 1 litre of mud in the Davis and wells. He concluded that Ingolfiella is either of and I. from coral lives in inaccessible Kara- Strait, east Greenland, littoralis, very rare or biotopes (S. sands on the shores ofThailand. Several later years man, 1959). anotherspecies, I. acherontis (S. Karaman, 1933), Meanwhile, another species was found in South was described from a groundwater well in Skopje, America in a new habitat, the coastal ground water Macedonia. The material ofI. acherontis has been in a coarse shingle beach. In the description of I. few lost and only a incomplete drawings remain. ruffoi Slewing, 1958, Slewing posed a dilemma S. Karaman apparently was not aware of the work particularly relevant to the work underlying this by Hansen, 1903. Therefore he describedBalcanella publication. He asked whether the large African acherontis in form the and n. gen. n. sp. a new family Balcanel- cave was ancestral to smaller marine lidae. freshwater if it the other around. forms, or was way After the Second World War expeditions focused Another type of habitat was added to the list more on environments that with the collection of of britannica were difficult to access, 20 specimens I. and the of majority species ofingolfiellideans were Spooner, I960, from the shell gravel at the sea collected in half the last of the twentieth century. bottom at 46 meter depth off the south coast of A in large species was found cave waters from England. Spooner remarked on the true deep sub- Downloaded from Brill.com10/10/2021 03:55:37AM via free access Contributions to Zoology, 72 (1) - 2003 41 surface occurrence of these ingolfiellids in contrast From the same environment, but a little deeper at in of with other small malacostracans that more typically 8 m below the surface the Gulf Naples, Italy, inhabit only the first few centimeters of these gravels. Schiecke (1976) described I. ischitana. Stock named tolerance revealed within Euryhaline was one 6 species from islands off the coast of Venezuela. I. manni species with the discovery of Noodt, 1961, These species are: I. (Gevgeliella) putealis Stock, taken from both brackish and fresh ground water 1976, in a slightly brackish well on Bonaire (Dutch in Chile. In addition, I. manni was found not only Antilles); I. (Gevgeliella) fontinalis Stock, 1977, at sea level, also at 800 m. Later I. but uspallatae from a spring on Bonaire; I. (Gevgeliella) tabularis 1965 was described from in the Noodt, Argentina Stock, 1977, from marine sands situated below a riverbanks of a at 2000 m in the Andes floodplain cave entrance on Bonaire (also recorded on Cura- mountains. cao); I. (Hansenliella?)quadridentata Stock, 1979, In at this two freshwater Europe time, species from coarse coral sand of a submarine flat, Curacao; were reported from the interstices of river alluvia I. (Trianguliella?) grandispina Stock, 1979, pumped of southern France; I. catalanensis Comeau, 1963 in from brackish ground water gravel at a cave and I. thibaudi Coineau, 1968. This marked the entrance; I. (Gevgeliella) margaritae Stock, 1979, of where subsur- discovery more species special for from a capped freshwater well used drinking face habitats were sampled in a conscious effort to water on Isla de Margarita (Venezuela). These south- A 2.3 uncover new taxa. large cm ingolfiellid was ern Caribbean species were sampled either by div- found in a well in first described as Leleu- Namibia, in shallow coastal from ing water, or pumped up piella eggerti, Ruffo, 1964, later renamed by Ruffo beach interstitia and oligohaline water reservoirs (1974a) as Trogloleleupia eggerti. more inland. In addition, animals occurred in wells From the Indian Ocean I. xarifae Ruffo, 1966 and small cave chambers with brackish water or and I. & kapuri Coineau Rao, 1973 were reported water with frequently and strongly fluctuating sa- from the Maldives in shallow coral sands, and from linities. the Andaman and Nicobar Islands in intertidal mud- Stock utilized subgeneric names, but the split- shell dy debris, respectively. ting of the genus Ingolfiella into smaller units had A second deep-sea species, I. atlantisi Mills, 1967, already begun with that earlier discovery of In- was collected 4700 at more than m depth in the golfiella (Balcanella) acherontis Karaman, 1933. North American Basin of the Atlantic Ocean. This Nevertheless, Karaman’s original effort did not I. species probably occupies the same habitat as useful due to subsequent workers’ lack abyssi prove very and was found amidst typical abyssal fauna of in male or female specimens cases where both elements, presumably residing “...in the flocculent sexes were needed to make critical distinctions. and relatively oxidized upper centimeters or two Consequently, new species were difficult to incor- on the surface of the deep-sea ooze.” (Mills, 1967). into the existing system. Back in porate Europe, an environment equally diffi- Ruffo (1985) described another large ingolfiellid cult to access as the deep sea was tapped, namely the from Namibia’s freshwater reservoirs, Stygobar- ‘fossil water’ of in a 50 m deep well Italy. Here The the nardia caprellinoides. most striking