Sissom, W . D . and O . F. Francke. 1983 .

Post-birth development of Vaejovis bilineatus Pocock

(Scorpions : Vaejovidae) . J . Arachnol ., 11 :69-75 .

POST-BIRTH DEVELOPMENT OF VAEJOVIS

BILINEATUS POCOCK (SCORPIONES : VAEJOVIDAE )

W. David Sissom' and Oscar F . Francke

Department of Biological Sciences

Texas Tech University
Lubbock, Texas 7940 9

ABSTRACT

The post-birth development of Vaejovis bilineatus Pocock was studied in the laboratory . Litter size
varied from 17 to 26 among five litters reared . The average growth factor between successive molts for
carapace length was 1 .26 ± 0 .05 ; for chela length 1 .28 ± 0 .05 ; and for metasomal segment V lengt h
1 .32 ± 0 .07 . Although no specimens were reared from birth to maturity, it was determined by extra-
polation and comparison with field-collected adults that there are six instars (five molts) to maturity .

INTRODUCTION

Only two studies dealing with post-birth development of the Vaejovidae have bee n
published to date . Francke (1977) reared Uroctous mordax Thorell in the laboratory ,
but only one male successfully reached the fifth instar . From its size at that age, Franck e
estimated that maturity was reached at the seventh instar . Polis and Farley (1979) cond-
ucted field studies on Paruroctonus mesaensis Stahnke . They reported that (1) maturity
was reached at 19 to 24 months of age, (2) there were seven instars in the post-birt h
development, and (3) growth was determinate . In addition they discussed ecologica l
parameters affecting the life history of P. mesaensis and presented a summary of al l
published scorpion life histories in tabular form .
In addition to these studies, there have been several others of related interest . McAl-
ister (1960) discussed growth rates of first instar young of Vaejovis spinigerus (Wood) .
Williams (1969) published information on birth behavior for Anuroctonus phaiodactylus
(Wood), Uroctonus mordax Thorell, Vaejovis confusus Stahnke, V. minimus Kraepelin ,
V. spinigerus, V. vorhiesi Stahnke, and P. mesaensis . He also included descriptions of firs t

and second instars of those species . Haradon (1972) published additional information o n
birth behavior for U. mordax . Finally, Hjelle (1974) published observations on the birth
and post-birth behaviors of Syntropis macrura Kraepelin .
The present study represents the first developmental information available for th e
genus Vaejovis beyond the second instar . Five pregnant females of Vaejovis bilineatus
Pocock were collected at Villa Hidalgo, San Luis Potosi, Mexico in March 1977 . They
gave birth in the laboratory in August and their litters (n = 17, 20, 22, 25, and 26) wer e

' Present Address : Department of General Biology ; Station B, Box 1812 ; Vanderbilt University ;
Nashville, Tennessee 37235 .


70 THE JOURNAL OF ARACHNOLOG Y

reared . As the litter of one female (n = 17) died as second instars, the specimens wer e
omitted from all calculations . Fourteen specimens from the other litters entered the
fourth instar and one of those entered the fifth . An immature female collected with th e
adults at Villa Hidalgo molted twice and reached maturity in the laboratory . The speci-
mens from these two sources provided a complete study of the post-birth development o f
V. bilineatus.

MATERIALS AND METHODS

Rearing methods used in this study have been described elsewhere (Francke 1977 ,
1979) . Briefly, young scorpions born in the laboratory were separated into individua l
containers as soon as the litters dispersed from the females' dorsa . A layer of soil wa s
provided as a substrate, and water was provided with a moistened sponge . Early instars
were fed macerated bits of roaches (Nauphoeta cinerea Saussure), as available living prey
were too large for the young scorpions to subdue . The later instars were given live imma-
ture N. cinerea . All specimens were maintained in an environmental chamber at near
constant temperature (26 .6 ± 2°C) .
Two problems resulted in loss of specimens and/or data . The macerated bits of roaches
used as food molded rapidly and young scorpions became trapped in the fungal hyphae .
The specimens were killed and usually damaged as a result . Other specimens died afte r
burrowing in the soil and dessicated so that measurements were useless .
Measurements of carapace length, pedipalp chela length, and metasomal segment V
length were taken for all instars observed (either from exuviae or preserved specimens) .
All measurements were taken with an American Optical Model 569 binocular dissectin g
microscope equipped with an ocular micrometer calibrated at 20X .
Growth factors between successive instars were calculated from the data by dividin g
the dimension of a structure at one instar by its dimension at the previous instar . An
average growth factor and the standard deviation were calculated for each molt from th e
pooled data . In the text and tables, all average ages, measurements, and growth factors ar e
accompanied by the standard deviation (in the form of x ± s) unless otherwise specified .
The failure of specimens to reach maturity required the use of extrapolation an d
calculation of 95% confidence intervals of instars not observed in the litters (see Franck e
1979 for procedure) . Measurements of field-collected adults from the same or nearb y
localities were compared to predicted size ranges of instars, and the instar at whic h
maturity is reached was thus inferred .
RESULTS

First Instar.—The young of Vaejovis bilineatus are born enclosed by a "birth mem-
brane " as are other apoikogenous scorpions (Francke, in press) . Upon freeing themselve s
from this membrane the young climb onto the mother's dorsum and orient themselves i n
rows with the prosoma directed forward and the metasoma backward . This layering effect
was reported by Williams (1969) for several Vaejovis spp . and is apparently characteristi c
of the genus . We have observed the same for V. coahuilae Williams and V. waueri Gertsch
and Soleglad .
The first molt occurred at an age of 9 .78 ± 0 .41 days . Molting of all the young took
place in a single day . Dispersion of the young took two to four days to complete, and this
process began at an age of 16 .8 ± 1 .7 days (7 .0 ± 1 .6 days after the first molt) .
First instars of V. bilineatus are creamy white at birth and gradually darken until time
of exuviation . As in all other scorpions studied to date, they lack many adult features .


SISSOM AND FRANCKE–POST-BIRTH DEVELOPMENT OF VAEJOVIS BILINEATUS 71

Tarsi are blunt and lack claws ; the aculeus is soft and blunt ; trichobothria, setae, granula-
tion, and carinae are absent ; and the denticles on the margins of the pedipalp chela finger s
are absent . Pectines, however, are well developed and contain the complete number o f
pectinal teeth .
Second Instar.—Laboratory mortality was very high in the second instar . Only 40 of
93 specimens (43 .0%) survived this stage, and most of the deaths were probably due t o
dessication either prior to or during molting . There is good evidence for mortality fro m
molting difficulties : specimens dying during or around peak molting periods had formed a
new cuticle which was visible underneath the semi-transparent second-instar cuticle in th e
carapace region .
Specimens surviving the second instar (n = 40) molted to the third instar at an age o f
106 .2 ± 15 .6 days . The duration of the second instar was 96 .2 ± 15 .7 days (range =
68-140 days) (Table 1) . Measurements of second instar structures and growth factors fo r
the second molt are found in Table 2 . Growth factor information could be obtained for
only 31 of the 40 specimens : seven specimens dessicated before measurements were take n
and two died during the second molt (there was no cuticular expansion and hardening) .
The average growth factor from second to third instar in carapace length was 1 .26 ± 0 .06 ;
in chela length 1 .28 ± 0 .05 ; and in metasoma V length 1 .33 ± 0 .07 .
Second instars possess most of the adult characters . The body is covered with setae ,
and all trichobothria are present . The tarsi bear claws, the aculeus is hardened and sharp ,
and the dentate margins of the pedipalp chela forgers are developed . Coloration is essen-
tially the same as in adults . Carinae of the pedipalps and metasoma are very poorl y
developed (carinal development is gradual throughout life) .
Third Instar .—Of the 40 specimens reaching the third instar, only 14 (35%) complete d
it . Specimens surviving this instar molted to the fourth instar at an age of 190 .1 ± 27 . 9
days . The duration of the third instar was 87 .5 ± 20 .4 days (range = 56-140 days) (Table
1) . Measurements of third instar structures and growth factors for the third molt ar e
found in Table 2 . Only nine of 14 specimens yielded growth factor information : three
specimens dessicated during the fourth instar (measurements were not possible) and tw o
died during the third molt . The average growth factor from third to fourth instar i n
carapace length was 1 .26 ± 0 .04 ; in chela length 1 .27 ± 0 .04 ; and in metasoma V lengt h
1 .30 ± 0 .04.
Fourth and Succeeding Instars .—All remaining young except one died during th e
fourth instar . The surviving specimen molted to the fifth instar at an age of 323 days ; th e
duration of its fourth instar was 147 days (Table 1) . Measurements of fourth and fift h
instar structures are reported in Table 2 .
One immature female was collected with the pregnant females at Villa Hidalgo ; i t
molted twice in the laboratory and attained maturity . Growth factors for this specimen
did not differ from the laboratory-reared litters (Table 2) . The duration of its penultimate
instar was 243 days . The specimen did not molt between January 1978 and its death i n
October 1981, so apparently there is no post-reproductive molt in this species . This is
further evidence that post-reproductive molts do not occur in the Scorpiones .
Using methods described by Francke (1979), 95% confidence intervals were calculate d
for the fifth, sixth, seventh, and eighth instars. These confidence intervals were calculate d
from measurements and growth factors of fourth instar specimens (n = 9), rather than
from the single fifth specimen . The confidence intervals for carapace length, chela length ,
and metasoma V length are found in Table 2 .


72 THE JOURNAL OF ARACHNOLOG Y

Table 1 .-Duration (Dur .) of instars and cumulative age (Cum .) at time of molting in days i n
laboratory-reared Vaejovis bilineatus Pocock . Only specimens surviving the second instar are shown .
Deaths occuring during a given instar are indicated by "x". Specimens lost are indicated by "?".
First instar means and standard deviations are based on all specimens from the four litters used (n =
93) . The number in parentheses following the specimen number (Spec . No .) indicates to which litter
the specimen belongs .

First Second Third Fourth

Spec. No . Dur. Cum . Dur . Cum . Dur. Cum. Dur. Cum.

1(1) 10 10 131 141 098 239 x

2(1) 10 10 100 110 093 203 x
3(1) 10 10 100 110 x
4(1) 10 10 086 096 x
5(1) 10 10 089 099 ?
6(1) 10 10 105 115 081 196 x
7(1) 10 10 086 096 x
8(1) 10 10 095 105 x
9(1) 10 10 073 083 x
10(1) 10 10 089 099 x
11(1) 10 10 088 098 090 188 x
12(1) 10 10 098 108 x
13(1) 10 10 080 090 x
14(1) 10 10 100 110 109 219 ?
15(1) 10 10 068 078 072 150 x
16(1) 10 10 098 108 x
17(1) 10 10 101 111 x
18(2) 9 9 093 102 140 242 x
19(2) 9 9 118 127 x
20(2) 9 9 083 092 101 19 3
21(2) 9 9 118 127 x
22(2) 9 9 090 099 x
23(2) 9 9 090 099 056 155 x
24(2) 9 9 123 132 x
25(3) 10 10 105 115 x
26(3) 10 10 092 102 x
27(3) 10 10 094 104 076 180 x
28(3) 10 10 072 082 x
29(3) 10 10 096 106 x
30(3) 10 10 102 112 x
31(3) 10 10 094 104 072 176 147 32 3
32(4) 10 10 103 113 x
33(4) 10 10 140 150 x
34(4) 10 10 079 089 082 171 x
35(4) 10 10 108 118 x
36(4) 10 10 070 080 x
37(4) 10 10 086 096 078 174 x
38(4) 10 10 100 110 x
39(4) 10 10 115 125 x
40(4) 10 10 089 099 077 176 x

n 93 40 40 14 14 1 1
9.78 96 .2 106 .2 87 .5 190.1 147 32 3
s 0.41 15 .7 15 .6 20 .4 27 .9 --


SISSOM AND FRANCKE—POST-BIRTH DEVELOPMENT OF VAEJOVIS BILINEATUS 73

Table 2 .—Measurements in mm and growth factors of observed instars (mean ± standard deviation) ;
predicted 95% confidence intervals for instars not observed in the laboratory ; and measurements of
field-collected specimens of Vaejovis bilineatus Pocock .
* denotes estimated instars of collected specimens .

Carapace Chela Metasoma V

OBSERVE D
2nd instar (n = 88) 1 .43 ± 0 .07 1 .61 ± 0 .08 1 .34 ± 0.0 9
Growth factor (n = 31) 1 .26 ± 0 .06 1 .28 ± 0 .05 1 .33 ± 0 .0 7
3rd instar (n = 31) 1 .86 ± 0 .13 2 .09 ± 0 .15 1 .82 ± 0 .1 7
Growth factor (n = 9) 1 .26 ± 0 .04 1 .27 ± 0 .04 1 .30 ± 0 .0 4
4th instar (n = 9) 2.30 ± 0 .13 2 .58 ± 0 .18 2 .30 ± 0.1 7
Growth factor (n = 1) 1 .23 1 .29 1 .3 6
5th instar (n = 1) 2 .74 3 .30 2 .8 9
Female molting in laborator y
4th instar* 2 .56 2.89 2 .6 1
Growth factor 1 .23 1 .26 1 .3 3
5th instar* 3 .15 3 .65 3 .4 6
Growth factor 1 .29 1 .34 1 .2 7
6th instar* 4 .05 4 .90 4 .4 0
PREDICTED 95% CONFIDENCE INTERVAL S
5th instar 2.57—3 .22 2 .85—3 .75 2 .54—3 .4 9
6th instar 3 .23 — 4 .06 3 .65 — 4 .80 3 .30 — 4 .5 4
7th instar 4 .07 — 5 .12 4 .67 — 6 .15 4 .29 — 5 .9 0
8th instar 5 .13 — 6 .45 5 .98 — 7 .87 5 .57 — 7 .6 6
FIELD-CAUGHT SPECIMEN S
Adult females (n = 9) 3 .83 ± 0 .39 4 .62 ± 0.30 4 .14 ± 0 .2 6
Adult males (n = 2 ; range) 2 .98 — 3 .25 3 .90 — 4 .10 3 .55 — 3 .7 5
4th instar* (n = 2 ; range) 2.40 — 2 .50 3 .00 2 .55 — 2 .6 0
5th instar* (n = 2 ; range) 2.75 — 2 .85 3 .30 — 3 .60 2 .90 — 3 .4 0

DISCUSSIO N

When measurements of field-collected adults are compared to the confidence interval s

obtained from the laboratory-reared litters (Table 2, Fig . 1), most of them (73%) fit
easily within the intervals predicted for the sixth instar . However, three adult female s
(two giving birth to laboratory-reared litters and the immature specimen molting twice i n
the laboratory) fall within the lower limits of the seventh instar . Although it is possible
that these three females are indeed seventh instars, we consider them to be large sixt h
instar adults based on the following observations . The young of one litter are significantly
larger than the others (Duncan's multiple range test ; X's = 1 .37, 1 .41, 1 .41, 1 .42, 1 .52 ; p
< 0 .01), and extrapolation from the average size of second instars (using the growt h
factor obtained from this litter) shows that a specimen of this litter could easily fit in th e
predicted seventh instar confidence intervals after only four molts . The fact that som e
specimens do not fit in the " correct " size range suggests limitations in the method, bu t
this is primarily due to the difficulty in rearing large numbers of scorpions past th e
early instars .
Only two adult males were available for comparison . In chela and metasoma V lengt h
both specimens fit the sixth instar confidence intervals . In carapace length one specime n
is slightly smaller than predicted ; the other is just within the lower limit for the sixth


74 THE JOURNAL OF ARACHNOLOGY

5 -

t
7p t

4
ti
•t
6p

3 I •
5p
I • •I

4o
r
2–
3o

► r ► , ,

2 3 4 5 6

CHELA LENGTH (MM )

Fig . 1 .–Plot of carapace length versus pedipalp chela length in instars of Vaejovis bilineatus . Boxes
with solid lines depict 95% confidence intervals for instars observed (0) in the laboratory (2nd throug h
4th) ; boxes with broken lines depict 95% confidence intervals for predicted (p) instars (5th throug h
7th) . Symbols are as follows : o =,,field-collected specimens ; A = exuviae and specimen of female molt-
ing to maturity in the laboratory .

instar . This could be attributable to sexual allometry, but sample sizes of both field -
collected adults and laboratory-reared specimens were too small to permit statistica l
analyses .
As stated previously there are potential problems with estimating size ranges of instar s
not observed . To minimize these problems, the estimations should be derived from large
sample sizes of late instars, as this reduces error considerably . Another problem encount-
ered in using this method is that of allometric growth at the maturation molt . This is
common in scorpions (e .g ., allometry in the metasomal segments in male Centruroides
spp .), but with prior knowledge of its occurrence, adjustments can be made with reason -
able accuracy (Sissom and Francke, unpublished data) .
Data for instar duration obtained in the laboratory probably do not reflect duration s
in the field, as feeding regimes certainly differ . These data are reported here as they ma y
prove useful in future laboratory studies .

ACKNOWLEDGMENTS

This study was completed in partial fulfillment of the requirements for the degree o f
Master of Science in the Department of Biological Sciences at Texas Tech University (W .


SISSOM AND FRANCKE—POST-BIRTH DEVELOPMENT OF VAEJOVIS BILINEATUS 75

D . Sissom) . We wish to thank the following individuals for help in collection or care of
specimens : James C . Cokendolpher, Steven K . Jones, Robert W . Mitchell, and Frederick
W . Wagner . We also thank Mr . Cokendolpher for his comments and criticism of th e
manuscript . A very special thanks to Lisa A . Nelms for her help and encouragement t o
the senior author during the course of the study. This study was supported in part by a
Graduate Student - Faculty Summer Fellowship from the Graduate School, Texas Tec h
University, and by the Institute for Museum Research .

LITERATURE CITE D

Francke, O . F . 1977 . Obervations on the life history of Uroctonus mordax Thorell (Scorpionida ,
Vaejovidae) . Bull. British Arachnol. Soc., 3(9) :254-260 .
Francke, O . F . 1979 . Observations on the reproductive biology and life history of Megacormus gert-
schi Diaz (Scorpiones : Chactidae : Megacorminae) . J . Arachnol ., 7 :223-230 .
Francke, O . F . In press . Parturition in scorpions (Arachnida, Scorpiones) : A review of the ideas. Rev.
Arachnol .
Haradon, R . M. 1972 . Birth behavior of the scorpion Uroctonus mordax Thorell (Vaejovidae) . Ent .
News, 83 :218-221 .
Hjelle, J . T . 1974 . Observations on the birth and post-birth behavior of Syntropis macrura Kraepelin
(Scorpionida : Vaejovidae) . J . Arachnol ., 1 :221-227 .
McAlister, W. H . 1960 . Early growth rates in offspring from two broods of Vejovis spinigerus Wood .
Texas J . Science, 4 :162-168 .
Polis, G . A . and R . D . Farley. 1979 . Characteristics and environmental determinants of natality ,
growth and maturity in a natural population of the desert scorpion, Paruroctonus mesaensis
(Scorpionida : Vaejovidae) . J . Zool ., London, 187 :517-542 .
Williams, S . C . 1969 Birth activities of some North American scorpions. Proc . California Acad . Sci. ,
37(1) :1-24 .

Manuscript received October 1981, revised January 1982.