Home Page for:

Denson McLain

Dept. of Biology
Georgia Southern University



Born in Baytown, Texas, 13 November 1953. I love my wife, cats, numbers, photography, amateur astronomy, snorkeling, and sailing. I like to sit by myself in my office, with Ann nearby, doing statistics and listening to Rush Limbaugh.



My primary research focus is the impact of the environment (physical, biotic, or social) on mate choice and sexual selection. Currently, my wife, Ann Pratt, my Ph.D. advisor, Donald J. Shure, and I have two study insect systems that we have now studied for over 30 years (work on each began in 1979). These are (1) mate choice and the evolution of ethological isolation in the soldier beetle, conducted every autumn in the foothills and mountains of northern Georgia (Cornelia, Helen, Hiawassee, Clayton), and (2) territorial behavior of the seed bug on the granite outcrop, Arabia-Davidson Mountain, near Lithonia, Georgia. Currently, we are addressing questions that require multi-year data sets. For instance, in the soldier beetle we are documenting the repeated appearance and disappearance of assortative mating by wing spot length (i.e., beetles with a locality behaving as two species for a few years then reverting to behaving as a single species). In the seed bug, we are looking at the impact of climate variability on the abundance of the bug and in turn how abundance affects mating dynamics and interspecific competition for host plant seeds. Ann and I continue to work on a variety of fiddler crabs species in the Virgin Islands, Bahamas, and along the Atlantic coast from the Florida Keys to Cape Cod. We are especially interested in factors that favor smaller or larger claws (the mass of the single male claw can be 60% of that of the rest of the body). We have identified habitat, density, and presence of competing fiddler crab species as important factors possibly affecting the development and level of selection on claw size and shape.

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My Curriculum Vitae

Emvironmental Biology - Practice Problems

THESE RULES AND FORMULAE WILL APPEAR ON EVERY EXAM.

Rules for entering numbers on the scan form.

Mark 1 or more letters on every scan form line allocated to an answer; each number or symbol in the answer is recorded on its own scan-form line, in order beginning with the leftmost number or symbol.

Begin all negative answers with [BCD] on the first answer line.

For answers with an absolute value of <1, do not put any 0’s to the left of the decimal point; Positive answers <1 begin with [ABCDE], negative answers begin with BCD on the first line and [ABCDE] on the second line allocated for the answer.

Do not round any answers.

If an answer is not an integer (whole number) enter as much of its fractional part (numbers to the right of the decimal point) as can be recorded in the lines allocated for the answer; record 0’s [A] on extra lines if the fractional part terminates.

If there are more answer lines than numbers in the answer, place a decimal [ABCDE] then 0’s [A], if necessary, on remaining answer lines.

Record an answer of 0 as [A] if there is only 1 line allocated or as [ABCDE] followed by [A]’s if more than 1 line is allocated.


Letters stand for numbers and mathematical symbols. Minus sign: [BCD] Decimal point: [ABCDE]

Numbers: [A] 0 [B] 1 [C] 2 [D] 3 [E] 4

[AB] 5 [AC] 6 [AD] 7 [AE] 8 [BC] 9

r = birth rate – death rate Nt = N0●er●t t1/2 = 70/(100●[-r])

tdouble = 70/(100●r) ln(x) = y; ey = x ln(ey) = y

eln(x) = x ln x = a; ea = x ln(ea●b) = a●b

NG = N0●(R0)G R0 = ∑(lx●mx) T = [∑(x●lx●mx)]/R0

r = [ln(R0)]/T vt = [∑(lx●mx)]/lt S = c•Az

ln(a●b) = ln(a) + ln(b) ln(ab) = b●ln(a)


THIS WILL NOT APPEAR ON EXAMS. Commit this to memory: r = rate of increase, b - d; b = birth rate = Births/Sample size; d = death rate = Deaths/Sample size; t = years; Nt = population size t years from now or t years from the beginning; N0 = population size 0 years from now (i.e., right now) or the original (beginning, initial) population size.

SAMPLE PROBLEMS FOR EXAM 1.

A random sample of 1250 females from a population of 150,000 was monitored for 1 year. Among the sample females, there were 65 births of a daughter and 50 deaths. Assume birth and death rates remain constant.

1-6. What is the birth rate?

Birth rate ≠ number of births; birth rate = number births χ number of females sampled (monitored), i.e., 65 χ 1250 (=65/1250) = .05200

Recorded as: line 1 – ABCDE
line 2 – A
line 3 – AB
line 4 – C
line 5 – A
line 6 – A

7-10. What is the death rate?

Death rate ≠ number of deaths; death rate = number deaths χ number of females sampled (monitored), i.e., 50 χ 1250 (=50/1250) = .040

Recorded as: line 7 – ABCDE
line 8 – A
line 9 – E
line 10 – A

11-15. What is the rate of increase?

Want to calculate r, which is the birth rate minus death rate; i.e. r = b-d = (65/1250) – (50/1250) = 15/1250 = .012

Recorded as: line 11 – ABCDE
line 12 – A
line 13 – B
line 14 – C

16-20. Using the rule of 70 formula, how long before the population doubles in size?

tdouble = 70/(100∙r) = 70χ[100∙(.012)] = 70/1.2 = 58.333

Recorded as: line 16 – AB
line 17 – AE
line 18 – ABCDE
line 19 – D
line 20 – D

21-25. Using the rule of 70 formula, how long before the population grows to 8X its current size?

tdouble = 58.333. How many times must the population double to increase by 8X?

Note the series: 1 → 2 → 4 → 8→ 16 → 32 → 64 → 128. Each arrow represents 1 doubling time. To increase from 1 to 8 requires doubling to 2, then doubling to 4, then doubling to 8, i.e., 3 doublings (3 arrows). The answer is the number of arrows (doublings) times the doubling time. The answer here is 3∙58.333 = 175.0 years.

Recorded as: line 21 – B
line 22 – AD
line 23 – AB
line 24 – ABCDE
line 25 – A

If the population was declining, this series,1→ 1/2 → 1/4 → 1/8→ 1/16 → 1/32 → 1/64 → 1/128, could be used in the same way with the t½ formula.

26-31. How large will the population be in 25 years?

Use the formula Nt = N0∙e r∙t. Nt = N25 = the population size in 25 years. N0 is the population size 0 years from now. In this set of populations, N0 is 150,000. Often, N0 is given in the original statement. Do not confuse N0 with the sample size (S), the number of females monitored or censused for 1 year (here, S = 1250). In the exponent for e, r is the rate of increase, 0.012, and t is the number of years through which the population will grow, 25. Using the ex button, key in “e^(.012 x 25)”. Multiply this answer by N0 (150,000).

Nt = N0∙e r∙t, or N25 = 150,000∙[e .012∙25] = 150,000∙[e .30] =150,000∙(1.349858808) = 202,478. Note: never round numbers, and carry as many significant digits as you can throughout all calculations.

Recorded as: line 26 – C
line 27 – A
line 28 – C
line 29 – E
line 30 – AD
line 31 – AE

32-33. How many years must pass before the population reaches 190,000?

Here, find “t” in the formula Nt = N0∙e r∙t. Nt is given, 190,000, and N0 was given in the original problem statement as equal to 150,000. The value of “r” has already been determined to be 0.012. Start by filling all known values into the equation, Nt = N0∙e r∙t. This gives:
190,000 = 150,000∙e 0.012 ∙ t. The key to solving for “t” (or “r” in a similar problem [see below]) is to eliminate the “e”. Remember, in algebra, a mathematical operation done to one side of an equation must be done to the other side. Begin by simplifying; i.e., divide each side by 150,000. This gives: 190,000/150,000 = [150,000/150,000]∙e 0.012 ∙ t → 1.2666… = e 0.012 ∙ t. Now, take the natural log (ln) of each side (because ln[ey] = y, and, here, y is r∙t). This gives
ln[1.2666…] = ln[e 0.012 ∙ t] which all simplifies to: 0.236388778 = 0.012∙t. Dividing each side by 0.012 gives 0.236388778 / 0.012 = [0.012 / 0.012]∙t → 19.69906 = t.

Recorded as: line 32 – B
line 33 – BC

34-40. What value of r would lead to a tripling of the population in 50 years?

Now, find “r” in the formula Nt = N0∙e r∙t. Nt is to be 3 times N0, so Nt = 450,000. N0 is still the value given in the original problem statement, 150,000. The value of “t” has been specified now to be 50 years. Again, Start by filling all known values into the equation, Nt = N0∙e r∙t. This gives: 450,000 = 150,000∙e r ∙ 50. The key to solving for “r” (as when solving for “t” in a similar problem [see above]) is to eliminate the “e”. Remember, in algebra, a mathematical operation done to one side of an equation must be done to the other side. Begin by simplifying; i.e., divide each side by 150,000. This gives: 450,000/150,000 = [150,000/150,000]∙e r ∙ 50 →
3 = e r ∙ 50. Now, take the natural log (ln) of each side (because ln[ey] = y, and, here, y is r∙t). This gives ln[3] = ln[e r ∙ 50t] which all simplifies to: 1.098612289 = r∙50. Dividing each side by 50 gives the value of r: 1.098612289 / 50 = [50 / 50]∙ r → r = .021972246

Recorded as: line 34 – ABCDE
line 35 – A
line 36 – C
line 37 – B
line 38 – BC
line 39 – AD
line 40 – C

r = b – d = (B – D)/S Nt = N0 • er•t t double or half = 70/(100•r), where


THIS WILL NOT APPEAR ON EXAMS. Commit this to memory: r = rate of increase; b = birth rate = Births/Sample size; d = death rate = Deaths/Sample size; t = years; Nt = population size t years from now; N0 = population size 0 years from now (i.e., right now). vx = reproductive value (number of daughters remaining to be born to the average female of age x); lt = survival from birth to age t (t = age now; value of lt does not change in vx equation); lX = survival from birth to age x (changes for each age across which sums, Σ, are made); mx = age-specific fecundity (production of daughters; changes for each age across which sums, Σ, are made); R0 = net reproductive rate; T = generation length; x = specific age (or middle of an age group); NG = population size in G generations of length T; N0 = population size 0 generations from now (i.e., current size); G = number of generations over which the population will grow.

PROBLEMS using DEMOGRAPHIC DATA. Use the information in the table below to answer questions 1-65. The demographic data refer to a Polynesian population just before the arrival of Europeans. The size of the population is 15,000.

Probability of survival from birth Age-specific fecundity Median age in years Age group (years)
1.00 0 .001 0-.003
0.95 0 4.5 .003-9
0.90 0.6 13.5 9.003-18
0.80 1.0 22.5 18.003-27
0.65 0.3 31.5 27.003-36
0.45 0.1 40.5 36.003-45
0.20 0.01 49.5 45.003-54
0.01 0 63.0 54.003-72
0.00 - - >72

Same data

Age group
0-.003
.003-9 9.003-18 18.003-27 27.003-36 36.003-45 45.003-54 54.003-72 >72

Age
0
4.5
13.5
22.5
31.5
30.5
49.5
63.0
-
Survival from birth
1.00
0.95
0.90
0.80
0.65
0.45
0.20
0.01
0.00
Age-specific fecundity
0.00
0.00
0.60
1.00
0.30
0.10
0.01
0.00
-

1. What is the net reproductive rate?

2. What is the length of a generation?

3. What is the reproductive value of a newborn female (in the 0-0.003 age group)?

4. What is the reproductive value of a female of age 18.003 years?

5. How large will the population be in 3 generations?

6. What is the rate of increase?

7. How large will the population be in 21 years.

8. What is the time (years) required for population size to double? (Use the rule of 70 formula.)

9. What is the time (years) required for the population to increase to 16 times its current size? (Use the rule of 70 formula.)

10. How many years must pass before the population reaches 25,000?

11. What would the generation length be if all age-specific fecundities decreased by 50%?

12. What would happen to generation length if only the 9.003-18 year old group saw its age-specific fecundity drop?
CHOICES: [A] decrease [B] increase [C] no change

13. What would happen to generation length if the survival rate increased for females passing from the 27.003-36 age group to the 36.003-45 age group?
CHOICES: [A] decrease [B] increase [C] no change

13. What would happen to generation length if only the 9.003-18 year old group saw its age--specific fecundity increase?
CHOICES: [A] decrease [B] increase [C] no change

Use these formulae.

r = birth rate – death rate Nt = N0●er●t t1/2 = 70/(100●[-r])

tdouble = 70/(100●r) ln(x) = y; ey = x ln(ey) = y

eln(x) = x ln x = a; ea = x ln(ea●b) = a●b

NG = N0●(R0)G R0 = ∑(lx●mx) T = [∑(x●lx●mx)]/R0

r = [ln(R0)]/T vt = (1/ lt)•[∑(lx●mx)]

1. R0 = ∑(lx●mx). R0 = net reproductive rate. “∑” means to sum a set of numbers, here from the age of first reproduction (here:13.5 years old) to the age of last reproduction (here: 49.5 years). lx= the probability of surviving from age birth to age x (x just stands for some age). mx= the average number of daughters born to females of age x.

R0 = ∑(lx●mx) = (0.9 x 0.6) + (0.8 x 1.0) + (0.65 x 0.3) + (0.45 x 0.1) + (0.2 x 0.01) = 1.582

2. T = [∑(x●lx●mx)]/R0. Now sum the products of age (x), survival from birth (lx), and age-specific fecundity (mx), again from the ages of first to last reproduction, then divide by the net reproductive rate.

T = [∑(x●lx●mx)]/R0 = [(13.5 x 0.9 x 0.6) + (22.5 x 0.8 x 1.0) + (31.5 x 0.65 x 0.3) + (40.5 x 0.45 x 0.1) + (49.5 x 0.2 x 0.01)] / 1.582 = 21.083

T will usually be close to the age when mx peaks (here: mx peaks at 22.5). If you calculate a value that is very different from the age where mx peaks, check for a mistake (like forgetting to divide by R0).

3. vt = [∑(lx●mx)]/lt. Reproductive value is the average number of daughters remaining to be born to females of a specified age. When the specified age, or age group, includes newly born females, vt = v0 = R0. The answer = 1.582.

4. vt = [∑(lx●mx)]/lt. Reproductive value is calculated similarly to the net reproductive rate except for 2 differences: (1) start summing the product, lx•mx, from the age specified, not from the age of first reproduction, and (2) after summing the products, divide by lt, which is just the probability of surviving from birth to the age specified (survival from birth to the age the female is right now). Dividing by lt credits the female for having survived over a period of time during which some females die (Note: lt for a newborn = 1.)

vt = [∑(lx●mx)]/lt = [(0.8 x 1.0) + (0.65 x 0.3) + (0.45 x 0.1) + (0.2 x 0.01)] / 0.8 = 1.3025

5. NG = N0●(R0)G, where N0 is the population size now (initial size) and G is the number of generations specified in the problem statement. (Note: G is not the same as T.)

N3 = N0●(R0)3 = 15000•(1.582 3) = 59,389

6. r = [ln(R0)]/T. r = the rate of increase (Note: r is not the same as R0. Pay attention to which parameter is used in a particular formula.)

r = [ln(R0)]/T = { ln(1.582) } / 21.083 = 0.021756

Do not forget to divide by T (generation length). When using T on an exam, do not divide by a rounded value.

7. Nt = N0●er●t. Nt = 15000●e0.021756●21 = 15000●e0.456884 = 23,687.

A common mistake is to use R0 instead of r in the formula. Use common sense to check your answers – here 21 years is about 1 generation so this answer must be smaller than that in problem 5 which involves growth over 3 generations. Here, we expect the answer to be close to NG = N0●(R0)G where G = 1 (N1 = 15,000•[1.582 1] = 23,730).

8. tdouble = 70/(100●r). Again, be careful (by reading the formula) to use r not R0.

tdouble = 70/(100●0.021756) = 32 years

9. 16x = 2 x 2 x 2 x 2 = 24. Take the exponent 4 and multiply by tdouble
= 4 x 32.175 = 128.7. The population must double, double again, double again, and double again to grow to 16X its current size. Each of the 4 periods of doubling takes the same amount of time.

10. Nt = N0●er●t. Solve for t.

25000 = 15000●e0.021756●t next, divide through by 15000

1.6667 = e0.021756●t next, take the natural log of each side

ln(1.6667) = 0.021756•t taking the log of e raised to an exponent yields the exponent

0.5108 = 0.021756•t → 0.5108/0.021756 = t → t = 23.4

Syllabus - Spring 2011

Environmental Biology (BIOL 1230 C; CRN 10701). Spring 2011. D K McLain.

Lecture: 2:30 PM – 3:45 PM, Monday & Wednesday. Room 1119 Biology Bldg.

Website: http://www.georgiasouthern.edu/cost/biology; click on bio.georgiasouthern.edu; faculty McLain

Office: Room 1102 D, Biology. Office hours: 11:00 AM – noon, T Th. Texts: none.

Tentative Sequence of Topics
Introduction, biodiversity, & modeling population growth
Climate, seasons, & incident solar radiation
Water
The niche
Coral reefs & reef fish diversity
Reproductive strategies
Modeling population growth with schedules of reproduction & death
Reproductive value & age structure
Hawaiian HIPPO
Biomes – Part 1: African savannah
Biomes – Part 2: Tropical rainforest
Aquatic Habitats – Part 1: Intertidal zone
Aquatic Habitats – Part 2: Estuaries & marshes
Climate change
Preserving biodiversity: island biogeography & metapopulation dynamics
The scientific method - Part 1: assumptions, probability, & experimentation
Sound science
The scientific method – Part 2: comparative biology
Ecosystem structure - Part 1: feeding relationships
Energy metabolism: work, thermodynamics, & the coupling of anabolism & catabolism
Ecosystem structure - Part 2: symbioses & competition
Environmental ethics

Exam dates & times: Exam 1 – Wednesday, 9 February; Exam 2 – Wednesday, 9 March; Exam 3 – Wednesday, 20 April; Exam 4 (Comprehensive Final Exam) – Monday, 9 May (3:00 PM – 5:00 PM). Students arriving after an exam has been passed out may not be permitted to take the exam.

Exam format: The following types of questions or prompts are likely – problem solving, true/false, matching, & multiple-choice. Answering problems accounts for about 50% of the value of each exam. All exams will be machine graded. Therefore, students must bring Accu-scan form ABF-2052 (available at the bookstore) and a #2 pencil. It is the student’s responsibility to carefully mark the scan form so that it can be machine read – superfluous marks, light marking, and incomplete erasure can cause some answers to be graded as incorrect. Scan forms must not be folded or crumpled, even at the edges, as this may make machine-grading impossible – a penalty of -25 points will be applied to each exam for which machine grading is not possible. BRING 2 SCANFORMS TO THE FINAL EXAM.

Grading Policy: No make-up exams will be given with the possible exception of the final exam (see Website for Course Policies). Each exam constitutes 25% of the final average. The score you receive on an exam is based on the % of exam value answered correctly + any in-class bonus points earned. Bonus points earned during in-class exercises can apply only to the up-coming exam. Bonus points earned as part of an exam will only be applied to that exam. The final letter grade associated with the average of 4 grades: A = 100-89.5, B = 79.5-89.49999, C = 69.5-79.49999, D = 59.5-69.49999, F<59.5. Students missing an exam with a legitimate excuse (see Course Policies) will have the final exam count as 2 exams. Anyone missing 2 or more scheduled exams will fail the course.

Materials needed: # 2 pencil(s), Texas Instruments TI-30XA calculator ($12.99 at the bookstore; learn to use the ex and ln x keys), accu-scan ABF-2052 forms. If you do not bring this calculator to an exam, you will not be permitted to take the exam. Calculators may not be shared among students.
Course goals: To learn how organisms (including humans) interact with and affect their environment; to learn how ecological principles apply to everyday life; to learn about environmental issues that result from human interaction with the environment; to understand and promote environmentally responsible behavior.

Learning outcomes: Demonstrate and understanding of ecology, including populations, communities, and ecosystems and the role of evolution in shaping various levels of ecology; understand major environmental problems and how they relate to biological concepts and principles; think critically about environmental problems; understand environmental problems that occur or are associated with everyday life; demonstrate scientific literacy skills; demonstrate an understanding of the scientific process, including designing and interpreting experiments.

Policies:

Attendance: You may not make up any in-class bonus assignments done on days you fail to attend.

Academic honesty: Consult the Student Conduct Code for definitions of and policies on cheating. Students found violating the conditions of academic honesty will receive an F in the course. Additionally, violations of academic honesty will be reported to the administration which may entail additional penalties.

Classroom Disruption: Students disrupting the classroom will be assessed penalties on an accelerating scale. Examples of disruptive behavior include having your cell phone go off in class, talking on the phone, text messaging during class, using a computer in class for any purpose not related to the class, repeatedly entering class late or departing early, talking in class after you or another student has been warned to be quiet, arguing with or complaining to or about the professor, threatening a faculty member, physical display of anger or violence directed towards students or faculty. Penalties are deductions from the final average as follows – 1st offense, -5 points, 2nd offense, -10 points, 3rd offense, -20 points, 4th offense, -40 points. Disruptive students may be asked to leave the classroom or may be removed by security personnel.

Changes and Modifications: While provisions of the syllabus are as accurate and complete as possible, the instructor reserves the right to change any provisions herein without actual notice if circumstances so warrant. Every effort will be made to keep students advised of such changes and information about such changes will be available at all times from the instructor. It is the responsibility of each student to know what changes, if any, have been made to the provisions of this syllabus and to successfully complete the requirements of this course.
See the Policies statement posted on the professor’s website for information regarding civil conduct, academic dishonesty (cheating), legitimate excuses for missing the final exam, responses to fire alarms, etc.

 

 


 

Curriculum Vitae

Denson K McLain

Personal

Business Address:
Department of Biology, Georgia Southern University, Statesboro, Ga. 30460-8042
Business Phone:
(912) 478-5480
E-mail address:
dk_mclain@georgiasouthern.edu

Education

Texas A & M University.
August 1972 - May 1976. B.S. (Biology). Summa cum laude.

University of Florida.
September 1976 - June 1978. M.S. (Zoology). Non-thesis degree.

Emory University.
September 1978 - June 1982. Ph.D. (Biology). Dissertation: Behavioral and evolutionary ecology of the seed bug, Neacoryphus bicrucis, a host plant specialist on pyrrolizidine alkaloid-bearing plants (D.J. Shure, advisor).

Emory University.
July 1982 - July 1983. Postdoctoral research associate, Department of Microbiology (J.R. Scott, advisor). Construction of P1-derived plasmid vectors, molecular genetic analysis of bacteriophage P1 heat-shock mutants.

University of Notre Dame.
August 1983 - November 1985. Postdoctoral research associate, Department of Biology (K. S. Rai, advisor). Mosquito systematics, evolution of satellite DNA, evolution of ethological isolation.

Centers for Disease Control & Emory University.
January 1986 - August 1987. Postdoctoral research associate, National Research Council of the National Academy of the Sciences (F.H. Collins [CDC] & V. Finnerty [Emory University], advisors). Mosquito systematics and population structure, molecular genetic characterization of rDNA.

Academic Positions

University of Notre Dame.
August 1983 - November 1985. Postdoctoral research associate, Department of Biology (K. S. Rai, advisor). Mosquito systematics, evolution of satellite DNA, evolution of ethological isolation.

Centers for Disease Control & Emory University.
January 1986 - August 1987. Postdoctoral research associate, National Research Council of the National Academy of the Sciences (F.H. Collins [CDC] & V. Finnerty [Emory University], advisors). Mosquito systematics and population structure, molecular genetic characterization of rDNA.

Georgia Southern University, Department of Biology. August 1987 - present.

Teaching Experience and Expertise

Animal Anatomy, laboratory (BIO 480, BIOL 5241)
Biological Macrophotography (BIOL 5539)
Biology of Organisms Laboratory (BIOL 3112)
Cell Structure and Function (BIO 370)*
Cellular & Molecular Biology (BIOL 2111, 2113)
Environmental Biology (BIOL 1230)
Evolution (BIO 465/665)
Evolutionary Ecology (BIOL 5533)
Evolutionary Ecology (Evolution & Ecology; BIOL 7090)*
General Biology I (BIOL 1130)
Genetics (BIO 472/672)
Graduate Seminar (BIO 7610)
Population Biology (BIO 474/674; BIOL 5544)*
Principles of Biology I Laboratory (BIOL 2108L)
Principles of Ecology (BIO 473)
Population & Ecological Genetics (BIO 599/799)
Population & Quantitative Genetics (BIOL 5134)*
Sex & Evolution (BIOL 5535)
Undergraduate research (BIOL 4890)
Zoology (BIO 281)

* Recent, standard assignments

Publications

McLain D K (1979) Terrestrial trail following by three species of predatory stink bugs. Florida Entomologist 62: 152-4.

_________ (1980) Relationships among ants, aphids and coccinelids on wild lettuce. Journal of Entomological Science 15: 417-8.

_________ (1980) Female choice and the adaptive significance of prolonged copulation in Nezara viridula. Psyche 87: 325-36.

_________ (1981) Sperm precedence and prolonged copulation in the southern green stinkbug, Nezara viridula. Journal of Entomological Science 16: 70-7.

_________ (1981) Numerical responses of Murgantia histrionica to concentrations of its host plant. Journal of Entomological Science 16: 257-60.

_________ (1981) Responses of predatory pentatomid bugs to the excrement of the predator Euthyrhynchus floridanus (Hemiptera: Pentatomidae). Journal of Entomological Science 16: 535-7.

_________ (1981) Resource partitioning by three species of hemipteran herbivores on the basis of host plant density. Oecologia 48: 414-7.

_________ (1981) Benefit/cost ratios in the evolution of altruistic behavior toward parents and full siblings. Evolutionary Theory 5: 227-32.

_________ (1981) Interspecific interference competition and mate choice in the soldier beetle, Chauliognathus pennsylvanicus. Behavioral Ecology & Sociobiology 9: 65-6.

_________ (1982) Behavioral and morphological correlates of male dominance and courtship persistence in the blister beetle, Epicauta pennsylvanica (Coleoptera: Meloidae). American Midland Naturalist 107: 396-403.

_________ (1982) Density dependant sexual selection and positive assortative mating in natural populations of the soldier beetle, Chauliognathus pennsylvanicus. Evolution 36: 1227-35.

_________ (1983) Ants, extrafloral nectaries and herbivory on the passion vine, Passiflora incarnata. American Midland Naturalist 110:433-9.

_________ (1984) Host Plant Morphology, speciation, and the economics of mate choice in the soldier beetle, Chauliognathus pennsylvanicus. Evolutionary Theory 7: 63-7.

_________ (1984) Coevolution: Mullerian mimicry between a plant bug (Miridae) and a seed bug (Lygaeidae) and the relationship between host plant choice and unpalatability. Oikos 43: 143-8.

_________ (1984) Host plant density and territorial behavior of the seed bug, Neacoryphus bicrucis (Hemiptera: Lygaeidae). Behavioral Ecology & Sociobiology 14: 181-7.

_________ (1985) Clinical variation in morphology and assortative mating in the soldier beetle, Chauliognathus pennsylvanicus (Coleoptera: Cantharidae). Biological Journal of the Linnean Society 25: 105-17.

_________ (1985) Male size, sperm competition, and the intensity of sexual selection in the southern green stink bug, Nezara viridula (Hemiptera: Pentatomidae). Annals of the Entomological Society of America 78: 86-9.

McLain D K, Rai K S & Rao P N (1985) Ethological divergence in allopatry and asymmetrical isolation in the South Pacific Aedes scutellaris subgroup. Evolution 39: 998-1008.

McLain D K & Shure D J (1985) Host plant toxins and unpalatability of Neacoryphus bicrucis. Ecological Entomology 10: 291-8.

McLain D K (1986) Resource patchiness and the intensity of sexual selection in a resource-defending polygynous insect species. Oikos 47: 19-25.

_________ (1986) Niche differentiation and the evolution of ecological isolation in a soldier beetle hybrid zone. Oikos 47:153-67.

_________ (1986) Null models and the "intensity" of sexual selection. Evolutionary Theory 8: 49-51

McLain D K & Rai K S (1986) Reinforcement for ethological isolation in the southeast Asian Aedes albopictus subgroup. Evolution 40: 1346-50.

McLain D K, Rai K S & Fraser M J (1986) Interspecific variation in the abundance of highly repeated DNA sequences in the Aedes scutellaris (Diptera: Culicidae) subgroup. Annals of the Entomological Society of America 79:784-91.

McLain D K (1987) Heritability of size, a sexually selected character, and the response to sexual selection in a natural population of the southern green stinkbug, Nezara viridula (Hemiptera: Pentatomidae). Heredity 59: 391-5.

McLain D K & Boromisa R D (1987) Stabilizing sexual selection and density-dependent correlates of copulatory success in the ambush bug, Phymata wolffii (Hemiptera: Reduviidae). American Midland Naturalist 117:94-102.

_________ (1987) Interrelationship between male choice, fighting ability, assortative mating and intensity of sexual selection for the milkweed longhorn beetle, Tetraopes tetaophthalmus (Coleoptera: Cerambycidae). Behavioral Ecology & Sociobiology 20: 239-46.

McLain D K, Rai K S & Fraser M J (1987) Intraspecific and interspecific variation in the sequence and abundance of highly repeated DNA among mosquitos of the Aedes albopictus subgroup. Heredity 58: 373-81.

McLain D K & Shure D J (1987) Pseudocompetition: interspecific displacement through misdirected courtship. Oikos 49: 291-6.

McLain D K (1988) Male mating preferences and assortative mating in the soldier beetle. Evolution 42: 729-35.

Black W C, McLain D K & Rai K S (1989) Patterns of variation in the rDNA cistron within and among world populations of the mosquito, Aedes albopictus (Skuse). Genetics 121: 539-50.

McLain D K (1989) Prolonged copulation as a post-insemination guarding tactic in a natural population of the ragwort seed bug. Animal Behaviour 38: 659-64.

McLain D K & Collins F H (1989) Structure of rDNA in the mosquito Anopheles gambiae and rDNA sequence variation within and between species of the Anopheles gambiae complex. Heredity 62: 233-42.

McLain D K, Collins F H & Brandling-Bennet A D (1989) Microgeographic variation in rDNA intergenic spacers of Anopheles gambiae in western Kenya. Heredity 62: 257-64.

McLain D K, Hancock M & Hope M (1990) Fitness effects of nonrandom mating in the ragwort seed bug, Neacoryphus bicrucis (Hemiptera: Lygaeidae). Ethology Ecology & Evolution 2: 253-62.

McLain D K, Lanier D & Marsh N (1990) Effects of female size, mate size and number of copulations on fecundity, fertility and longevity in the southern -green stink bug, Nezara viridula (Hemiptera: Pentatomidae). Annals of the Entomological Society of America 83: 1130-6.

McLain D K & Marsh N B (1990) Individual sex ratio adjustment in response to the operational sex ratio in the southern green stink bug. Evolution 44: 1018-25.

_________ (1990) Male copulatory success: heritability and relationship to mate fecundity in the southern green stink bug, Nezara viridula (Hemiptera: Pentatomidae). Heredity 64: 161-7.

McLain D K, Marsh N B, Lopez J R & Drawdy J A (1990) Intravernal changes in the level of parasitization of the southern green stink bug, Nezara viridula (Hemiptera: Pentatomidae), by the feather-legged fly, Trichopoda pennipes: host sex, mating status, and body size as correlated factors. Journal of Entomological Science 25: 501-9.

McLain D K & Shure D J (1990) Spatial and temporal density dependence of host plant patch use by the ragwort seed bug, Neacoryphus bicrucis (Hemiptera: Lygaeidae). Oikos 52: 306-12.

McLain D K (1991) Heritability of size: a positive correlate of multiple fitness components in the southern green stink bug. Annals of the Entomological Society of America 84: 174-8.

_________ (1991) The r-K continuum and the relative effectiveness of sexual selection. Oikos 53: 263-5.

_________ (1991) Components of variance in male lifetime copulatory and reproductive success in a seed bug. Behavioral Ecology & Sociobiology 29: 121-6.

McLain D K & Mallard S D (1991) Sources and adaptive consequences of egg size variation in Nezara viridula (Hemiptera: Pentatomidae). Psyche 98: 135-64.

McLain DK (1992) Preference for polyandry in Nezara viridula (Hemiptera: Pentatomidae). Journal of Insect Behavior 5: 403-10.

_________ (1992) Population density and the intensity of sexual selection on body length in spatially or temporally restricted natural populations of a seed bug. Behavioral Ecology & Sociobiology 30: 347-56.

_________ (1992) Oviposition site preferences in Neacoryphus bicrucis (Hemiptera: Lygaeidae): responses to the density and dispersion of a single host plant species. Journal of Insect Behavior 5: 729-39.

Oliver J H Jr, Owsley M R, Hutcheson H J, James A M, Chen C, Irby W S, Dodson E M & McLain D K (1993) Conspecificity of the ticks Ixodes scapularis and I. dammini (Acari: Ixodidae). Journal of Medical Entomology 30: 54-63.

McLain D K, Burnette L & Deeds D (1993) Within season variation in the intensity of sexual selection on body size in the bug Margus obscurator (Hemiptera: Coreidae). Ethology Ecology & Evolution 5: 75-86.

McLain D K (1993) Cope's rules, sexual selection, and the loss of ecological plasticity. Oikos 68: 490-500.

Wesson M, McLain D K, Oliver J H, Piesman J & Collins F H (1993) Investigation of the validity of species status of Ixodes dammini (Acari: Ixodidae) using ribosomal DNA. Proceedings of the National Academy of Science (USA) 90: 10221-5.

McLain D K, Wesson D M, Oliver J H Jr & Collins F H (1995) Variation in rDNA ITS 1 among eastern populations of Ixodes scapularis (Acari: Ixodidae). Journal of Medical Entomology 32: 353-360.

McLain D K, Wesson D M, Oliver J H, Jr & Collins F H (1995) Evolution of the rDNA spacer, ITS 2, in the ticks Ixodes scapularis and I. pacificus (Acari: Ixodidae). Heredity 75: 303-19.

McLain D K, Moulton M P & Redfern T P (1995) Sexual selection and the risk of extinction of introduced birds on oceanic islands. Oikos 74: 27-34.

McLain D K (1998) Direct benefits of mate choice: fecundity enhancement and sexy sons. Animal Behaviour 55: 1191-1201.

McLain D K & Vives S P (1998) Sexual selection and community structure: an island biogeographic analysis with beetles. Oikos 82: 271-281.

McLain D K, Moulton M P & Sanderson J G (1999) Sexual selection and extinction: the fate of plumage-dimorphic and plumage-monomorphic birds introduced onto islands. Evolutionary Ecology Research 1: 549-565.

McLain D K & Pratt A E (1999) Nestedness of obligate coral reef fish across a set of fringing reefs. Oikos 85: 53-67.

McLain D K & Pratt A E (1999) The cost of sexual coercion and heterospecific sexual harassment on the fecundity of a host-specific, seed-eating insect. Behavioral Ecology & Sociobiology 46: 164-170.

McLain D K, Moulton M P, Setters D L & Pratt A E. (2000) Ascription of resemblance of newborns by parents and nonrelatives. Evolution & Human Behavior 21: 11-23.

McLain D K, Li J & Oliver JH Jr (2001) Interspecific and geographic variation in the sequence of rDNA expansion segment D3 of Ixodes Ticks (Acari: Ixodidae). Heredity 86: 234-242.

McLain D K (2001) Evolution of transcript structure and base composition of rDNA expansion segment D3 of ticks. Heredity 87: 544-557.

Pratt A E, McLain D K & Kirschstein K (2002) Intrageneric predation by fiddler crabs in South Carolina. Journal of Crustacean Biology 22: 1-10.

Pratt A E & McLain D K (2002) Antisymmetry in male fiddler crabs and the decision to feed or breed. Functional Ecology 16: 89-98.

Pratt A E, McLain D K & Lathrop G R (2003) Sequential assessment in sand fiddler crab contests for breeding burrows. Animal Behaviour 69: 945-955.

McLain D K, Pratt A E & Berry A (2003) Predation by red-jointed fiddler crabs on congeners: interaction between body size and positive allometry of the sexually-selected claw. Behavioral Ecology 14: 741-747.

Mixson T R, Fang Q Q, McLain D K & Oliver J H Jr. (2004) Population structure of the blacklegged tick Ixodes scapularis revealed by SSCP data using the mitochondrial Cyt b and the nuclear ITS1 markers. Acta Zoologica Sinica 50: 176-186.

McLain D K, Pratt A E & Berry A (2005) Adaptive variation in the boldness of courting sand fiddler crabs (Uca pugilator ). Ethology 111: 63-76.

McLain D K, Pratt A E & Kirschstein K (2005) Predator-driven fragmentation of fiddler crab droves into mini-selfish herds of biased composition. Journal of Experimental Marine Biology & Ecology 315: 1-15.

McLain D K (2005) Female soldier beetles display a flexible preference for selectively favored male phenotypes. Evolution 59: 1085-1095.

Pratt A E & McLain D K (2006) How dear is my enemy: intruder-resident and resident-resident encounters in male sand fiddler crabs (Uca pugilator). Behaviour 143: 597-617.

McLain D K & Pratt A E (2007) Approach of females to magnified reflections indicates that claw size of waving fiddler crabs correlates with signaling effectiveness. Journal of Experimental Marine Biology and Ecology 343: 227-238.

McLain D K & Pratt A E (2008) Asymmetry of leg size and differential leg usage in the sand fiddler crab, Uca Pugilator. Journal of Crustacean Biology 28: 601-606.

Weese D A, McLain D K, Pratt A E, & Fang Q Q (2009) Population structure of the Atlantic fiddler crab (Uca pugilator) along the eastern coast of the US revealed by molecular data. Current Zoology 2: 150-157.

Moulton, M. P., McLain, D. K., & Moulton, L. E. 2009. Sexual selection and the fate of introduced pigeons and doves (Columbidae). Evolutionary Ecology Research 11: 200-208.

McLain, D. K., McBrayer, L. D., Pratt, A. E., & Moore, S. 2010. Performance capacity of fiddler crab males with regenerated versus original claws and success by claw type in territorial contests. Ethology, Ecology, & Evolution 22: 36-48.

McLain, D. K. & Pratt, A. E. 2010. Available energy in beach and marsh habitats and the size of the fiddler crab claw, a sexually selected weapon and signal. Oikos 119:508-513.

McLain, D. K. & Pratt, A. E. 2011. Body and claw size at autotomy affect the morphology of regenerated claws of the sand fiddler crab, Uca pugilator. Journal of Crustacean Biology (in press).

Papers presented

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Last Updated on 1/31/11 by Denson McLain.



Department of Biology, Georgia Southern University

last modified: 1/31/11