Fiddler Crabs Home

Unusual Development

The following is a collection of references to unusual development of secondary sexual characters in fiddler crabs, divided by basic type. It is not meant bo be a thorough overview of developmental studies. This also does not include anomalies in claw size due to regeneration status.

Males with two large claws (a.k.a.Super-males)

The most commonly observed unusual development is the rare finding of a male with two large claws, rather than the typical one large and one small. The reason this is the most commonly observed is not necessarily that it is the most common in nature, just that it is the easiest to identify as this strange quirk of morphology can readily be seen through binoculars (unlike the others, which all require more direct examination of the reproductive organs). Males with two large claws are presumed to be at a disadvantage with respect to obtaining food, as feeding is generally done with the small claws, but as individuals with this condition are occasionally found in nature, they clearly can find a way to survive. There is some suggestion that this state is seen more commonly in laboratory-reared individuals where there may be reduced selection pressure.

Double-clawed males
A pair of double-clawed male fiddler crabs of the species Minuca rapax, raised in an aquarium (Photo from H.-G. Rupp).

A secondary issue is some of these studies have focused on very young juveniles, where the asymmetry may not be entirely set. It is not uncommon to find tiny crabs with two larger claws where these later transition to standard asymmetry as the further develop. This is very different than the rarer instances of full-sized adult males with two large claws.

Smith (1870) Minuca pugnax One found in nature
Morgan (1920) Leptuca pugilator One found in nature
Morgan (1923) Minuca pugnax (Juvenile study) Reports 56 very small crabs (<3mm) found in nature with two large claws.
Vernberg & Costlow (1966) Minuca rapax (Juvenile study) Four males found in a lab reared population.
Takeda & Yamaguchi (1973) Gelasimus jocelynae One found in nature
Gibbs (1974) Minuca burgersi One found in nature
Williams & Heng (1981) Gelasimus vomeris One found in nature
Mulstay (1987) Minuca pugnax One found in nature
Monserrat & Rodríguez (1995) Leptuca uruguayensis One found in nature
Yamaguchi (1977) Austruca lactea (Juvenile study) Observed 300 lab-reared juveniles (out of ~14,000) with two large claws.
Yamaguchi (2001) Austruca lactea Mentions a few instances seen in young laboratory-reared crabs.
Yamaguchi & Henmi (2001) Tubuca arcuata One found in nature. Additional small juveniles with two large claws were observed, but experimentally this was shown to be a transient state that would not be (normally) maintained through adulthood.
Benetti & Negreiros-Fransozo (2003) Minuca panema Two found in nature
Lira et al. (2006) Leptuca cumulanta One found in nature
Odhano et al. (2015) Tubuca alcocki One found in nature
Vale et al. (2015) Minuca rapax One found in nature
iNaturalist Observation Minuca minax One found in nature
iNaturalist Observation Minuca minax One found in nature
iNaturalist Observation Tubuca arcuata One found in nature
iNaturalist Observation Leptuca pugilator One found in nature; species ID not 100%
iNaturalist Observation Paraleptuca splendida One found in nature
iNaturalist Observation Leptuca pugilator One found in nature; species ID not 100%
iNaturalist Observation Leptuca pugilator One found in nature
iNaturalist Observation Leptuca pugilator One found in nature
iNaturalist Observation Austruca mjoebergi One found in nature

Males with two small claws

Occasionally when examining large populations of fiddler crabs, male crabs have been found with two small claws largely indistinguishable from those of the females. Males with this condition would be at no survival disadvantage with respect to feeding, but might presumably have difficulty attracting mates. For very small juvenile male fiddler crabs, this is the standard condition before asymmetry develops. Males and females are largely indistinguisable at very small sizes as both asymmetry and other secondary sexual characteristics have not yet developed.

Morgan (1920) Minuca pugnax Six found in nature; in some one of the claws was slightly larger than the other, but overall still much more resembled a female in size and shape
Morgan (1920) Leptuca pugilator One found in nature
Takeda & Yamaguchi (1973) Gelasimus jocelynae Two found in nature
Ahmed (1976) Leptuca cumulanta Five young adults found in nature, although see comment below.
Benetti & Negreiros-Fransozo (2003) Minuca panema Four found in nature

Females with one large and one small claw

A single instance of an otherwise fully functional female crab (based on exmination and dissection of the ovaries) with a single large claw and a small claw (otherwise identical to those in male fiddlers) has been reported in the literature. As with the previous case, this condition would not be obvious in wild populations as the crab would otherwise superficially appear male in the field.

Deecaraman & Milton (1992) Leptuca pugilator One found in a shipment from a laboratory supply company

Intersex crabs

Intersex crabs are those with a seeming mix of male and female features, generally such that the crab is neither clearly male nor female, even upon examination of the reproductive organs. This is a slightly different (although arguably overlapping) category than the previous two in which the reproductive organs appeared to be unequivocally male or female, with only the asymmetry of the claws reversed from normal.

Morgan (1920) Minuca pugnax Reports 16 individuals with two small claws, but abdomens in between in size between males and females and often with unclear additional characteristics.
Morgan (1920) Leptuca pugilator Reports 8 individuals with two small claws, but abdomens in between in size between males and females and often with unclear additional characteristics.
Holthuis (1959) Minuca rapax Crab found in nature with two overly large claws (a bit smaller than those of a standard male), with an abdomen intermediate in size between male and female, and with pleopods similar to females, but with the exopod underdeveloped in the first two.
Ahmed (1976) Leptuca cumulanta In addition to the males with small claws listed above, he reports a number of what otherwise appeared to be female crabs with intermediate sized abdomens lacking pleopods. This study was focused on very young crabs and may in some cases represent delayed development in juveniles, rather than true intersex crabs.
Zou & Fingerman (2000) Leptuca pugilator One found in a shipment from a laboratory supply company. Had reduced bilateral asymmetry in the claws, an abdomen in size between male and female, and a mix of both male and female pleopods on different segments.